Frequency of Cyanogenesis in Tropical Rainforests of Far ...

28
Frequency of Cyanogenesis in Tropical Rainforests of Far North Queensland, Australia REBECCA E. MILLER*,RIGEL JENSEN and IAN E. WOODROW School of Botany, The University of Melbourne, Victoria, 3010, Australia Received: 14 January 2005 Returned for revision: 21 December 2005 Accepted: 23 January 2006 Published electronically: 6 March 2006 Background and Aims Plant cyanogenesis is the release of toxic cyanide from endogenous cyanide-containing compounds, typically cyanogenic glycosides. Despite a large body of phytochemical, taxonomic and ecological work on cyanogenic species, little is known of their frequency in natural plant communities. This study aimed to investigate the frequency of cyanogenesis in Australian tropical rainforests. Secondary aims were to quantify the cyanogenic glycoside content of tissues, to investigate intra-plant and intra-population variation in cyanogenic glycoside concentration and to appraise the potential chemotaxonomic significance of any findings in relation to the distribution of cyanogenesis in related taxa. Methods All species in six 200 m 2 plots at each of five sites across lowland, upland and highland tropical rainforest were screened for cyanogenesis using Feigl–Anger indicator papers. The concentrations of cyanogenic glycosides were accurately determined for all cyanogenic individuals. Key Results Over 400 species from 87 plant families were screened. Overall, 18 species (45 %) were cyanogenic, accounting for 73 % of total stem basal area. Cyanogenesis has not previously been reported for 17 of the 18 species, 13 of which are endemic to Australia. Several species belong to plant families or orders in which cyanogenesis has been little reported, if at all (e.g. Elaeocarpaceae, Myrsinaceae, Araliaceae and Lamiaceae). A number of species contained concentrations of cyanogenic glycosides among the highest ever reported for mature leaves—up to 52 mg CN g 1 d. wt, for example, in leaves of Elaeocarpus sericopetalus. There was significant variation in cyano- genic glycoside concentration within individuals; young leaves and reproductive tissues typically had higher cyanogen content. In addition, there was substantial variation in cyanogenic glycoside content within populations of single species. Conclusions This study expands the limited knowledge of the frequency of cyanogenesis in natural plant communities, includes novel reports of cyanogenesis among a range of taxa and characterizes patterns in intra-plant and intra-population variation of cyanogensis. Key words: Australia, b-glycosidase, chemotaxonomy, cyanogenic glycoside, cyanogenesis, defence, hydrogen cyanide, polymorphism, Queensland, screening, secondary metabolite, tropical rainforest. INTRODUCTION Cyanogenesis is the ability to release toxic hydrogen cyanide (HCN) from endogenous cyanide-containing com- pounds. It has long been recognized in plants (Conn, 1991; Seigler, 1991), and has been recorded in ferns, fern-allies, gymnosperms, as well as monocotyledonous and dicotyle- donous angiosperms from >550 genera and 130 plant families (Conn, 1981; Poulton, 1990; Jones, 1998). Cyano- genesis in plants requires the presence of either an unstable cyanohydrin, or of a stable cyanogen and its degradative enzymes (Seigler, 1991). While cyanolipids have been identified from a few taxa (Mikolajczak et al., 1970), cyano- genesis in plants most commonly results from the hydrolysis of cyanogenic glycosides (Conn, 1981). Autotoxicity in intact plants is prevented by the spatial separation— either at the subcellular or at the tissue level—of the cyano- genic glycoside and catabolic enzymes (Kojima et al., 1979; Selmar, 1993a; Poulton and Li, 1994; Zheng and Poulton, 1995; Hickel et al., 1996). The catabolism of cyanogenic glycosides is therefore initiated upon tissue disruption, due to mechanical damage or ingestion by herbivores, for example, which enables mixing of enzymes and cyanogenic substrate (Wajant et al., 1994; Patton et al., 1997). Little is known about the frequency of cyanogenesis in natural plant communities. This is despite a large body of literature documenting cyanogenesis in >2650 species of angiosperms worldwide (Lechtenberg and Nahrstedt, 1999). Indeed, as many as 11 % of all plant species are predicted to be cyanogenic (Jones, 1998). Historically, much of the interest in cyanogenesis centred around recording toxic plants with the potential for stock and human poisoning, the high frequency of cyanogenesis among food plants (Jones, 1998), and the potential utility of cyanogenesis and the structure of specific cyanogens in elucidating phylo- genetic relationships between taxa (e.g. Gibbs, 1974). As a consequence, much of what is known about the frequency of cyanogenesis comes from surveys of regional floras, or of specific taxonomic groups. There are a number of substantial chemotaxonomic works incorporating informa- tion on cyanogenesis (see Hegnauer, 1966, 1973, 1986, 1989, 1990; Gibbs, 1974), several smaller and more specific chemotaxonomic works (e.g. Tjon Sie Fat, 1978, 1979b; Spencer and Seigler, 1985; van Wyk and Whitehead, 1990; Seigler, 1994), including work on Australian Acacia spp. (Conn et al., 1985; Maslin et al., 1988), and numerous inventories of cyanogenic species (e.g. Rosenthaler, 1919; Seigler, 1976a, 1976b; Tjon Sie Fat, 1979a; Francisco and Pinotti, 2000). Several Australian researchers were active in * For correspondence. E-mail [email protected] Annals of Botany 97: 1017–1044, 2006 doi:10.1093/aob/mcl048, available online at www.aob.oxfordjournals.org Ó The Author 2006. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. 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Transcript of Frequency of Cyanogenesis in Tropical Rainforests of Far ...

Frequency of Cyanogenesis in Tropical Rainforests ofFar North Queensland Australia

REBECCA E MILLER RIGEL JENSEN and IAN E WOODROW

School of Botany The University of Melbourne Victoria 3010 Australia

Received 14 January 2005 Returned for revision 21 December 2005 Accepted 23 January 2006 Published electronically 6 March 2006

Background and Aims Plant cyanogenesis is the release of toxic cyanide from endogenous cyanide-containingcompounds typically cyanogenic glycosides Despite a large body of phytochemical taxonomic and ecologicalwork on cyanogenic species little is known of their frequency in natural plant communities This study aimed toinvestigate the frequency of cyanogenesis in Australian tropical rainforests Secondary aims were to quantify thecyanogenic glycoside content of tissues to investigate intra-plant and intra-population variation in cyanogenicglycoside concentration and to appraise the potential chemotaxonomic significance of any findings in relation to thedistribution of cyanogenesis in related taxa Methods All species in six 200m2 plots at each of five sites across lowland upland and highland tropical rainforestwere screened for cyanogenesis using FeiglndashAnger indicator papers The concentrations of cyanogenic glycosideswere accurately determined for all cyanogenic individuals Key Results Over 400 species from 87 plant families were screened Overall 18 species (45) were cyanogenicaccounting for 73 of total stem basal area Cyanogenesis has not previously been reported for 17 of the 18 species13 of which are endemic to Australia Several species belong to plant families or orders in which cyanogenesis hasbeen little reported if at all (eg Elaeocarpaceae Myrsinaceae Araliaceae and Lamiaceae) A number of speciescontained concentrations of cyanogenic glycosides among the highest ever reported for mature leavesmdashup to52mgCNg1 d wt for example in leaves of Elaeocarpus sericopetalus There was significant variation in cyano-genic glycoside concentration within individuals young leaves and reproductive tissues typically had highercyanogen content In addition there was substantial variation in cyanogenic glycoside content within populationsof single species Conclusions This study expands the limited knowledge of the frequency of cyanogenesis in natural plantcommunities includes novel reports of cyanogenesis among a range of taxa and characterizes patterns in intra-plantand intra-population variation of cyanogensis

Key words Australia b-glycosidase chemotaxonomy cyanogenic glycoside cyanogenesis defence hydrogen cyanidepolymorphism Queensland screening secondary metabolite tropical rainforest

INTRODUCTION

Cyanogenesis is the ability to release toxic hydrogencyanide (HCN) from endogenous cyanide-containing com-pounds It has long been recognized in plants (Conn 1991Seigler 1991) and has been recorded in ferns fern-alliesgymnosperms as well as monocotyledonous and dicotyle-donous angiosperms from gt550 genera and 130 plantfamilies (Conn 1981 Poulton 1990 Jones 1998) Cyano-genesis in plants requires the presence of either an unstablecyanohydrin or of a stable cyanogen and its degradativeenzymes (Seigler 1991) While cyanolipids have beenidentified from a few taxa (Mikolajczak et al 1970) cyano-genesis in plants most commonly results from the hydrolysisof cyanogenic glycosides (Conn 1981) Autotoxicity inintact plants is prevented by the spatial separationmdasheither at the subcellular or at the tissue levelmdashof the cyano-genic glycoside and catabolic enzymes (Kojima et al 1979Selmar 1993a Poulton and Li 1994 Zheng and Poulton1995 Hickel et al 1996) The catabolism of cyanogenicglycosides is therefore initiated upon tissue disruptiondue to mechanical damage or ingestion by herbivores forexample which enables mixing of enzymes and cyanogenicsubstrate (Wajant et al 1994 Patton et al 1997)

Little is known about the frequency of cyanogenesisin natural plant communities This is despite a large bodyof literature documenting cyanogenesis in gt2650 species ofangiosperms worldwide (Lechtenberg and Nahrstedt 1999)Indeed as many as 11 of all plant species are predicted tobe cyanogenic (Jones 1998) Historically much of theinterest in cyanogenesis centred around recording toxicplants with the potential for stock and human poisoningthe high frequency of cyanogenesis among food plants(Jones 1998) and the potential utility of cyanogenesisand the structure of specific cyanogens in elucidating phylo-genetic relationships between taxa (eg Gibbs 1974) As aconsequence much of what is known about the frequencyof cyanogenesis comes from surveys of regional floras orof specific taxonomic groups There are a number ofsubstantial chemotaxonomic works incorporating informa-tion on cyanogenesis (see Hegnauer 1966 1973 19861989 1990 Gibbs 1974) several smaller and more specificchemotaxonomic works (eg Tjon Sie Fat 1978 1979bSpencer and Seigler 1985 van Wyk and Whitehead1990 Seigler 1994) including work on Australian Acaciaspp (Conn et al 1985 Maslin et al 1988) and numerousinventories of cyanogenic species (eg Rosenthaler 1919Seigler 1976a 1976b Tjon Sie Fat 1979a Francisco andPinotti 2000) Several Australian researchers were active in For correspondence E-mail remunimelbeduau

Annals of Botany 97 1017ndash1044 2006

doi101093aobmcl048 available online at wwwaoboxfordjournalsorg

The Author 2006 Published by Oxford University Press on behalf of the Annals of Botany Company All rights reserved

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the field early last century reporting a number ofcyanogenic species and the specific cyanogenic consti-tuents involved (eg Petrie 1912 1913 1914 1920Finnemore and Cox 1928 1929 Finnemore andCooper 1936 1938) More than 700 species in theQueensland flora were screened by Smith and White(1918) Further phytochemical screening of the Queenslandflora was conducted by Webb (1948 1949) however therewas negligible testing among tropical taxa Importantlymany of these records are based on qualitative tests per-formed using herbarium specimens and tests of this kindusing dried material are by no means conclusive One fur-ther limitation of these data in terms of extrapolating tonatural plant communities is that negative results wereoften not reported

Tropical rainforests are of particular interest in thestudy of plant chemical defences Elevated herbivore pres-sure in tropical environments is hypothesized to havefavoured both a diverse array of defences and high levelsof investment in chemical defence (Levin 1976 Levin andYork 1978 Coley and Barone 1996 Kursar and Coley2003) Indeed in the field of plant secondary chemistry thetropical rainforest has been the focus of intense interestin co-evolutionary relationships between plants andherbivores and the extraordinary inter-specific and intra-plant variation in chemical defence strategies (Feeny1976 Levin 1976 Coley et al 1985 Coley andKursar 1996)

On the whole community-level studies of the distributionof chemical defences have focused on a small set of Asianand African rainforests (McKey et al 1978 Gartlan et al1980 Davies et al 1988 Waterman et al 1988) Inaddition these and other studies have tended to focus onC-based lsquoquantitativersquo defences (eg Coley 1983 1988)Among N-based defences the greater awareness of thefrequency of alkaloid-bearing plants in tropical and otherecosystems is a consequence of extensive phytochemicalscreening for bioactive compounds (eg Swanholm et al1960 Hartley et al 1973 Smolenski et al 1974 Collinset al 1990 Hadi and Bremmer 2001) or ecologicalresearch into specific plantndashherbivore interactions (egJanzen and Waterman 1984 Hartmann et al 1997) ratherthan systematic studies within natural communities (butsee Mali and Borges 2003)

There are some hypothesis-driven surveys for cyanogen-esis Two studies have investigated the frequency of cyano-genesis in floras investigating evolutionary and ecologicalhypotheses about exposure to herbivory (see Kaplan et al1983 Adsersen et al 1988 Adsersen and Adsersen 1993)However only the survey of Thomsen and Brimer (1997) inCosta Rican rainforest has been conducted in a systematicfashion well defined with respect to forest area and plantsize As with other N-based defences work on cyanogenictropical rainforest species worldwide is limited andthere has been no work on cyanogenesis in Australian trop-ical rainforests or any other form of chemical defence Thelack of work on cyanogenesis in diverse tropical rainforestsis further surprising as it is a readily detectable and con-stitutive chemical defence (Gleadow and Woodrow2000b)

Here we report some of the findings of a large-scalequantitative survey for cyanogenesis in Australian tropicalrainforests First we investigated the frequency of cyano-genesis and the contribution of cyanogenic species tobiomass (basal area) in lowland upland and highland trop-ical rainforest in north Queensland Australia Conductingthe survey in a standardized fashion will enable comparisonwith other communities (eg Thomsen and Brimer 1997)Secondly intra-plant and intra-population variation in con-centrations of cyanogenic glycosides was quantified Thehigh level of endemism among the Australian tropical rain-forest flora (Webb and Tracey 1981) and the number ofrainforest taxa previously untested for cyanogenesis under-score the potential for novel reports of cyanogenesis withindifferent taxonomic groups Finally therefore this studyaimed to investigate the potential taxonomic significanceof cyanogenesis reported here Overall this study aimedto expand the limited knowledge of the frequency ofcyanogenesis in the Australian flora and in natural plantcommunities

MATERIALS AND METHODS

Field sites

Field work was conducted between July 1999 andSeptember 2002 Six 200m2 plots (20 middot 10m) were estab-lished at each of five sites (total area 1200m2 per site) inlowland and upland rainforest in the tropics of north eastQueensland Australia (Fig 1) Six plots were selected fortwo reasons first the number of new species captured witheach additional plot had reached a plateau at around fivespecies and secondly it was a realistic sample size giventhe time and resources available Sites were selected tocapture maximum species diversity as on the AthertonTablelands forest type and species composition varyboth with substrate and with altitude (Tracey 1982) Fur-ther to enable comparison of forests occurring on differentsoil types two pairs of sites with similar altitude and rainfallwere selected The first pair comprised a site on soil derivedfrom basalt at Lamins Hill and one on soil derived fromgranite at Mt Nomico (Fig 1) The second pair comprisedsites on soils derived from basalt and rhyolite at LonglandsGap (Fig 1) A fifth site in lowland rainforest nearCape Tribulation and Myall Creek on soil derivedfrom metamorphic substrate was also surveyed (Fig 1)The distribution of cyanogenic species in relation toresource availability (soil nutrients) will be addressedelsewhere (see Miller 2004)

It was not possible to control strictly for logging historyall sites on the Atherton Tablelands had been selectivelylogged prior to the declaration of the Wet Tropics WorldHeritage Area in 1988 Detailed site descriptions are pro-vided in Miller (2004)

All individuals (palms trees and vines) with a diameter atbreast height (dbh) of gt5 cm were tagged and identifiedAll additional species (dbh lt5 cm) present in lower stratawith the exception of herbaceous ground species were alsotagged and identified

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Samples of all cyanogenic species as well as speciesdominant at each site and some rare species were pressedand lodged at the University of Melbourne (MELU) andBrisbane (BRI) Herbaria Accession numbers whereassigned are listed (Appendix)

Climate

The climate in far north Queensland is characterizedby a marked wet season from December to April Climaterecording stations in the study area are scattered thereforedata specific to each site were not available While locatedin the tropical latitudes because of its higher altitude theclimate of the Atherton Tableland is semi-tropical Meanannual temperature within the study area on the Tablelandis 22 C (Nix 1991) with a minimum of 10 C (Hall et al1981 see Graham et al 1995) All the upland and highlandrainforests surveyed on the Atherton Tableland havehigh average annual rainfall generally in the range2000ndash3000mm plus cloud interception (Tracey 1982)For example the average annual rainfall at Lamins Hillis 3584mm based on 30 years of records from the Queens-land Bureau of Meteorology (see Osunkoya et al 1993) Incontrast the climate in the coastal lowland tropical rain-forest near Cape Tribulation is characterized by highertemperatures Mean daily temperatures range from 28 Cin January to 22 C in July and temperatures may reach

the mid to high 30 s during the summer months Averageannual rainfall is also high at 3928mm recorded at CapeTribulation (based on 65 years of records from theQueensland Bureau of Meteorology)

Sites

Upland and highland rainforest The upland rainforest atLamins Hill [172240S 1454250E altitude 850m abovesea level (asl) Fig 1] is classified as complex mesophyllvine forest on basalt (type 1b Tracey 1982 Tracey andWebb 1975) This forest type typically occurs on uplandsites (400ndash800m) on high fertility kraznozem soils derivedfrom basalt with high rainfall (Tracey 1982) It is charac-terized by a closed canopy with multiple tree layers andan uneven canopy with height ranging from 35 to 45m(Tracey 1982)

The upland rainforest at Mt Nomico (171330S1454040E 900m asl Fig 1) is within the GilliesRange State Forest and is on low nutrient soil derivedfrom granite The forest is classified as complex notophyllvine forest typical of upland granitic soils (type 6 Tracey1982 Tracey and Webb 1975)

In the highland rainforest of Longlands Gap State Forest(altitude 1100ndash1200m asl) there is a sharp boundaryin forest type defined by basalt (172770S 1452850E)and rhyolite (172730S 1452860E) parent substrates

Atherton

AthertonTableland

MalandaLonglands Gap

0 50 km

Mareeba

Cairns

Port Douglas

Daintree River

Cape TribulationMyall Creek

WesternAustralia

SouthAustralia

NorthernTerritory

Queensland

New SouthWales

Lamins HillMt Nomico

F I G 1 Location of study sites in tropical rainforest in far north east Queensland Australia There were two pairs of sites in upland rainforest on the AthertonTablelandLaminsHill (172240S 1454250E) andMtNomico (171330S 1454040E) and two sites atLonglandsGap (17270S 145280E)Afifth sitewas

in lowland rainforest near Myall Creek and Cape Tribulation (160620S 1452690E)

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1019

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On basalt the forest is complex notophyll vine forest(type 5a Tracey 1982 Tracey and Webb 1975) a foresttype characterized by an uneven canopy (20ndash40m high) andnumerous tree layers typical of basaltic cool wet uplandsand highlands (Tracey 1982) On rhyolite the forest issimple microphyll vine forest (type 9 Tracey 1982)which is common on upland granitic soils (800ndash1300m)and is characterized by an uneven canopy 20ndash25m highwith emergent Agathis atropurpurea (up to 35m)

Lowland rainforest The fifth site was in lowlandrainforest in the Daintree World Heritage Area nearCape Tribulation (Fig 1) The site was near to Thompsonand Myall Creeks (160620S 1452690E altitude 40masl) The forest is complex mesophyll vine forest (type 1aTracey 1982) the canopy is irregular from 25 to 33m inheight and supports a great diversity of species and lifeforms including many palms and lianas The floristiccomposition is patchy with considerable variation incanopy and understorey dominants over small distances(Webb et al 1972 Tracey 1982) The soil is relativelynutrient-poor red clay loam podsol derived frommetamorphic substrate

Sampling for detection of cyanogenesis

Whole leaf samples (1ndash2 g f wt) were taken from indi-viduals with a dbh gt5 cm and from all additional speciesin the lower strata until at least three individuals of eachspecies had been sampled Where possible the youngestfully expanded leaves without epiphyllous communitieswere selected however in the case of samples acquiredby pruning shears attached to extension poles or by sling-shot and rope from the canopy it was not always possibleto be selective In instances where it was not possible toobtain samples from large canopy trees samples were takenfrom nearby individuals of the same species Fresh wholeleaf samples used for cyanide testing were stored in air-tightbags on ice until tested for cyanide 2ndash6 h later using FeiglndashAnger (FA) papers (Feigl and Anger 1966) Individuals ofrare species and those with little foliage were sampled onlyonce for analysis in the laboratory where a quantitativeassay was used to test for cyanogenesis using freeze-dried ground tissue Because cyanogenesis is known tobe a polymorphic trait (eg Hughes 1991 Aikman et al1996) a minimum of three individuals of all species wastested more where species were common Owing to thediverse and heterogeneous nature of the forests multipleindividuals of each species were not always represented inthe plots Therefore for these rare species testing indivi-duals located outside the plots was required and still therewere several species for which only single samples wereobtained

A range of other factors was taken into considerationwhen sampling For example given that young leavesof tropical species in particular are typically more highlydefended than older leaves (Coley 1983 Coley and Barone1996) both young (soft expanding leaves) and old (recentfully expanded) leaves were tested for cyanogenesis wherepossible In addition depending on availability fruit andflowers from several species were tested Owing to the large

number of species and samples in the survey it was notpossible to examine seasonal trends in defence howeverindividuals of the majority of species were tested in wetand dry seasons for qualitative changes in cyanogenic sta-tus as there is some evidence for seasonal variation incyanogenesis (Seigler 1976b Janzen et al 1980 Kaplanet al 1983) Finally because edaphic factors have beenreported to affect the expression of cyanogenesis (egUrbanska 1982) species found on more than one substratetype were tested at each site

Sample collection handling and storage for quantitativeanalysis

In addition to samples for fresh cyanide tests whole leafsamples (again recent fully expanded leaves) were taken forquantification of cyanogenic glycosides All individuals ofspecies that produced a positive result and individuals ofrare species were sampled Depending on sampling condi-tions samples were either placed immediately into liquidnitrogen or placed in a sealed air-tight bag and kept on icefor 2ndash6 h until snap frozen in liquid nitrogen Frozensamples were transported to the laboratory on dry icefreeze-dried and stored on desiccant at 20 C for analysisFreeze-dried samples were ground using either a cooledIKA Labortechnic A10 Analytical Mill (Janke and KunkelStanfen Germany) or for smaller samples an Ultramat 2Dental Grinder (Southern Dental Industries Ltd BayswaterVictoria Australia)

Chemical analyses

Detection of cyanogenesis FeiglndashAnger papers Thepresence of cyanogenic compounds in fresh field sampleswas determined using FA indicator papers (Feigl andAnger 1966) FA papers were selected in preference topicrate papers because FA papers are more sensitive(Nahrstedt 1980) and less prone to giving false-positiveresults (Brinker and Seigler 1989) FA test papers wereprepared according to Brinker and Seigler (1989) BecauseFA papers can be sensitive to moisture and light (Brinkerand Seigler 1989) they were stored in the dark and ondesiccant until use

Fresh leaves (approx 1ndash2 g f wt) were crushed induplicate screw-top vials Old and young foliage sampleswere tested separately To facilitate cyanogenesis 05mL ofwater was added to one of the vials and pectinase fromRhizopus spp (Macerase Pectinase 441201 CalbiochemCalbiochem-Novabiochem Corp La Jolla CA USA)(04 g Lndash1) in 01 M TrisndashHCl (pH 68) was added to theother Pectinase has been found to have non-specific b-glyc-osidase activity (Brimer et al 1995) and therefore inthe absence of sufficient endogenous b-glycosidase enablestests for the presence of cyanogenic glycosides to be madeThe indicator papers were suspended above the tissue bymeans of the screw-top lid and vials were left at roomtemperature and checked after 12 and 24 h This 24 htime period used in other surveys (eg Dickenmann1982 Thomsen and Brimer 1997 Buhrmester et al2000 Lewis and Zona 2000) was selected to avoid spuri-ous test results due to substantial bacterial contamination

1020 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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which may occur beyond 24 h (Saupe et al 1982) Tissue ofknown cyanogenic species Prunus turneriana or Ryparosajavanica was used as a positive control Moderate tostrongly cyanogenic samples gave a positive result withina few minutes or up to a few hours while more weaklycyanogenic samples took several more hours In accordancewith the recommendations of Brinker and Seigler (1989) anew test was conducted for any samples producing a slowpositive response (24 h) in case of interference by microbialcyanogenesis In addition any samples with inconclusivecolour change were re-tested An individual was consideredcyanogenic if a positive repeatable result was obtainedand a species was considered cyanogenic if at leastone individual produced a consistent and repeatablepositive result A negative test result indicates the absenceof a cyanogenic glycoside or of the specific cyanogenicb-glycosidase or both

In the few instances where insufficient tissue was avail-able for both FA paper tests using fresh leaves and sub-sequent laboratory analysis cyanogenesis was determinedbased on the quantitative assay of freeze-dried groundleaf tissue (as described below) by comparison with anegative tissue control (eg Alstonia scholaris or Aglaiameridionalis)

In this study tests for cyanogenesis used approx1ndash2 g f wt of leaves which is larger than tissue samplestested in previous surveys (eg 50mg fwt by Lewis andZona 2000 and 200mg f wt by Thomsen and Brimer1997 Buhrmester et al 2000) According to Dickenmann(1982) who used 500mg f wt tissue a weak positive reac-tion with FA papers where part of the paper turns blueindicated approx 2ndash20mgHCNkgndash1 f wt while a strongreaction indicated gt50mgHCNkgndash1 f wt These lowervalues equate to just over 6ndash60mgHCNgndash1 d wt using aconversion based on the mean foliar water content ofseveral species in this study which was 70 In thisstudy based on fresh leaf tests and quantitative analysisof freeze-dried tissue from the same sample the thresholdsensitivity of FA papers was similar within the range5ndash8mgHCNgndash1 d wt This threshold sensitivity of FApapers corresponds well to the criteria of Adsersenet al (1988) for classifying individuals as cyanogenicwhere individuals with lt93 nmol HCNgndash1 f wt (approxim-ately equivalent to 8mgHCNgndash1 d wt) were consideredacyanogenic

Polymorphism and confirmation of acyanogenesis

For the purposes of the survey a species was consideredcyanogenic if at least one individual produced a repeatableand positive test result For a few species not all individualsproduced positive results using FA papers therefore somefurther investigation of the polymorphism for cyanogenesisin these species was conducted First to confirm the acy-anogenic character of the individuals producing negativeresults with FA papers a greater mass of freeze-dried tissue(100mg) was assayed quantitatively for the evolution ofcyanide (see below) and compared with a non-cyanogenicspecies A scholaris as a control for baseline noise in theassay Based on the criteria of Adsersen et al (1988) if

lt8mg HCNgndash1 d wt was detected then that individual wasconsidered acyanogenic

Quantification of cyanogenic glycosides

The concentration of cyanogenic glycoside in plant tis-sues was measured by trapping the cyanide (CN) liberatedfollowing hydrolysis of the glycoside in a well containing1 MNaOH (Brinker and Seigler 1989 Gleadow et al1998) Freeze-dried ground plant tissue (30ndash50mg) wasincubated for 20ndash24 h at 37 C with 1mL of 01M citratendashHCl (pH 55) Cyanide in the NaOH well was determinedusing the method of Gleadow and Woodrow (2002) adaptedfrom Brinker and Seigler (1989) for use with a photometricmicroplate reader (Labsystems Multiskan Ascent withincubator Labsystems Helsinki Finland) The method ishighly sensitive the determination of CN in NaOH isrelatively specific for cyanide and can detect as little as5mg Lndash1 (Brinker and Seigler 1989) The absorbance wasmeasured at 590 nm with NaCN as the standard

RESULTS

Survey for cyanogenesis

In total fresh leaf material from 401 woody plant speciesfrom 87 families was tested for cyanogenesis (Appendix)Cyanogenesis was found in 18 species from 13 familiesrepresenting 45 of species occurring within 1200m2 ofrainforest at each of five sites In each case the addition ofpectinase during incubation did not result in any additionalpositive results The test with added enzyme thereforeeffectively served as a replicate test for each sample Afurther two species were found to be cyanogenic butwere not included in the analysismdashthe non-woody Alocasiabrisbanensis and Helicia nortoniana which occurred out-side the plots (Table 1) The number of cyanogenic speciesat each site ranged from four to 10 The cyanogenic speciesranged in life form from climbing monocotyledons suchas the supplejack Flagellaria indica to 30m canopytrees such as the northern silky oak Cardwellia sublimisThe proportion of cyanogenic species ranged from 47 inupland rainforest at Mt Nomico to 65 in lowland rain-forest near Cape Tribulation Overall cyanogenic speciesaccounted for 73 of total basal area (range 12ndash134)The highest proportion was in highland rainforest on basaltsoil at Longlands Gap while forest on rhyolite in adjacentforest had the lowest proportion In lowland rainforestthe contribution of cyanogenic species to biomass wasalso high at 116 compared with 33 and 71 at MtNomico and Lamins Hill sites respectively

The cyanogenic species and concentrations of cyano-genic glycosides in leaves and other plant parts aresummarized in Table 1 The foliar concentrations of cyano-genic glycosides among species ranged from around8mgCNgndash1 d wtmdashthe concentration below which individu-als were considered acyanogenic based on the criteria ofAdsersen et al (1988) and the limits of FA paperdetectionmdashto very high concentrations in excess of severalmgCNgndash1 d wt (Table 1) For example fully expanded

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1021

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TABLE1

Su

mm

ary

of

the

con

cen

tra

tio

no

fcy

an

og

enic

gly

cosi

des

inle

ave

sa

nd

oth

erp

lan

tti

ssu

esfr

om

cya

no

gen

icsp

ecie

sfo

un

din

six

plo

tsin

up

lan

dh

igh

land

(U)

an

dlo

wla

nd

(L)

tro

pic

alr

ain

fore

sta

tfive

site

sh

igh

nu

trie

ntb

asa

ltsi

tes

atL

am

ins

Hil

l(B

1)

an

dL

on

gla

nd

sG

ap

(B2

)a

nd

low

nu

trie

nts

ites

on

gra

nit

ea

tMtN

om

ico

(G)

on

rhyo

lite

at

Lo

ng

lan

ds

Ga

p(R

)a

nd

on

met

am

orp

hic

sub

stra

ten

ear

Ca

pe

Tri

bu

lati

on

(M)

Concentrationofcyanogenic

glycosides

indifferentplantparts(m

gCNg1dw

t)

Form

Species

Fam

ily

Forest

Site

Leaf

Stem

Floralparts

Fruitseed

HA

loca

sia

bri

sba

nen

sis

(FM

Bailey)Domin

Araceae

UB1

ndashndash

ndashndash

TB

eils

chm

ied

iaco

llin

aBHyland

Lauraceae

UB1B2GR

Old28 4ndash1263(n

=36)

ndashndash

ndashYg1039ndash1391

TB

rom

bya

pla

tyn

emaFM

uell

Rutaceae

LM

156 4ndash1285(n

=20)

ndashndash

ndash0ndash10 5

(n=27)

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

UL

B1B1GRM

Old11 5ndash69 8

(n=21)

ndashBud779ndash851

Seedapprox8

Yg733 1tips219 1

Flwr130ndash334

TC

leis

tanth

us

myr

ian

thu

s(H

assk)Kurz

Euphorbiaceae

LM

Old1 1ndash17 3

(n=41)

ndashFlwr30 7

Mature160ndash377

Yg78 4ndash80 4

(n=3)

Immature821

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

UB1B2G

Old1325ndash4800(n

=6)

ndashBud733

ndashFlwr440ndash942

TE

laeo

carp

us

seri

cop

etalu

sFM

uell

Elaeocarpaceae

UGR

1100ndash5056(n

=10)

Sdlg1739ndash2679

ndash1028

Tree

135

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

UB1B2

10ndash81(n

=3)

ndashndash

ndashV

Fla

gel

lari

ain

dic

aL

Flagellariaceae

UL

B1GM

11ndash177 3

(n=6)

ndashndash

ndashT

Hel

icia

aust

rala

sica

FM

uell

Proteaceae

LM

42 2ndash155 2

(n=8)dagger

ndashndash

ndashT

Hel

icia

bla

keiForeman

Proteaceae

UB1

Mean17 9

(n=2)

ndashndash

ndashT

Hel

icia

no

rto

nia

na

(FM

Bailey)FM

Bailey

Proteaceae

LM

ndashndash

ndashndash

VP

ass

iflo

rasp(K

urandaBH12896)

Passifloraceae

LM

313ndash922(n

=4)

ndashndash

ndashT

Mis

cho

carp

us

exa

ng

ula

tus

(FM

uell)Radlk

Sapindaceae

UB1

Old133yg222(n

=1)

ndashndash

ndash

TM

isch

oca

rpu

sg

ran

dis

sim

us

(FM

uell)Radlk

Sapindaceae

UL

GM

49ndash680(n

=2)

761ndash2006(n

=4)z

ndashndash

ndash

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

UB1B2

Old10ndash208(n

=7)

ndashndash

ndashYg2000

VP

ars

on

sia

lati

foli

a(Benth)

STBlake

Apocynaceae

UB1GR

765ndash4835(n

=3)

ndashndash

ndash

TP

oly

scia

sa

ust

rali

an

a(FM

uell)Philipson

Araliaceae

UL

B1B2GRM

Oldlt6

8(n

=30)10ndash28 5

(n=16)

ndashndash

ndashYg10 8ndash29 6tips259

TP

runu

stu

rner

ian

a(FM

Bailey)Kalkman

Rosaceae

UL

B1M

old2472ndash4888(n

=4)

560ndash1100(n

=2)

ndashMature

seed400

Yg3128ndash6464Tip6000ndash8498

Rootapprox2000

Ygfruitseed8950

TR

ypa

rosa

java

nic

a(Blume)

Kurz

exKoordamp

Valeton

Achariaceae

LM

1749(n

=2)

ndashndash

Mature

seed5430

Yglf2560(n

=5)

Ygseed7760(n

=5)

Meansandranges

ofcyanogenicglycosides

concentrationaregiven

forfullyexpanded

(old)leaveswherenotspecifiedandforsomeyoung(yg)leavesfor

nindividualsLifeform

herb(H

)tree

(T)vine(V

)Cyanogenic

glycosideconcentrationsweremeasuredas

evolved

cyanidefollowinghydrolysisoffreeze-dried

tissuewithouttheadditionofnon-specificb-glycosidase(egem

ulsinpectinase)

A

loca

sia

bri

sba

nen

sis(a

herb)and

Hel

icia

nort

onia

nawhichoccurred

outsidetheplotswerenotincluded

intheanalysisnorsampledforquantitativeanalysis

Concentrationsofcyanogenic

glycosides

inseedflower

andyoungleaves

of

Ryp

aro

saja

van

icadetermined

byBLWebber

(unpubldata2004Webber

andWoodrow2004)

daggerOnly

oneindividual

of

Hel

icia

aust

rala

sica

occurred

within

theplotsseven

additional

saplingstreeswereopportunisticallytested

externalto

theplotsallwerecyanogenic

zOnly

oneindividual

of

Mis

cho

carp

us

gra

nd

issi

mu

soccurred

insideplotsat

MtNomicoadditional

individualsweresampledoutsideplotsincludingfourin

lowlandrainforestallwerecyanogenic

1022 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

leaves of numerous individuals of Polysciasaustraliana tested negative for cyanide and containedlt8mgCNgndash1 d wt while others tested positive and werein the range of 10ndash28 mgCNgndash1 d wt Even in the weaklycyanogenic samples only in very few cases did the colora-tion of the test paper change from 12 to 24 h in a qualit-ative sense Individuals of numerous species had highconcentrations of cyanogenic glycosides in excess of1mgCNgndash1 d wt (eg Mischocarpus grandissimusBrombya platynema and Beilschmiedia collina) and severalspecies contained extremely high concentrations of cyano-genic glycosides in mature tree leaves Notably fullyexpanded leaves from the tree species Elaeocarpus serico-petalus Clerodendrum grayi and Prunus turneriana hadconcentrations of cyanogenic glycosides up to 52 49and 48mgCNgndash1 d wt respectively (Table 1)

Qualitative and quantitative variation in cyanogenesis

Intra-population variation in cyanogenic glycosides Forthe majority of species initially identified as being cyano-genic all individuals tested were cyanogenic however theconcentrations of cyanogenic glycosides varied markedlybetween conspecific individuals (Table 1) For example allsaplings and trees of B collina at each site tested positivefor cyanogenesis but the concentration of cyanogenic glyc-osides in fully expanded leaves from saplings (n = 19)ranged from 232 to 12634mgCNgndash1 d wt at Mt Nomicoalone where B collina was most abundant In contrast tothis quantitative phenotypic variation in cyanogenesis con-specific individuals of a small number of species producedboth positive and negative FA paper test results Thisapparent qualitative polymorphism was most notable inB platynema a subcanopy tree species restricted to thelowland rainforest Within the population of B platynemaapprox 50 of individuals tested produced negativeFA test paper results for both young and old leaves andhad cyanogenic glycoside concentrations in the range06ndash68mgCNgndash1 d wt (Table 1) The addition ofbuffered pectinase during testing did not alter the qualitativeFA paper test result for any individual Among cyanogenicB platynema the concentrations of cyanogenic glyco-sides varied over orders of magnitude from 105 to12859mgCN gndash1 d wt (Table 1)

Negative results with FA papers were obtained for twoother species Cleistanthus myrianthus and Polysciasaustraliana Qualitative and quantitative analysis of oldleaves from individuals of these species indicated a highfrequency of acyanogenic individuals however unlikeB platynema the phenotypic expression of cyanogenesisvaried qualitatively with leaf age such that young leavesand leaf tips from these same individuals were consistentlycyanogenic aswere reproductive tissues fromC myrianthus(Table 1)

Even among less common species with smallsample sizes the concentrations of cyanogenic glycosidesvaried markedly between individuals For example inMischocarpus grandissimus the concentration amongsix individuals from two sites ranged from 49 to680mgCNgndash1 d wt at Mt Nomico and from 637 to

2006mgCNg1 d wt in lowland rainforest and amongthree individuals of the woody vine Parsonsia latifoliaconcentrations ranged from 765 to 4835 mgCNg1 d wtThere was no detectable affect of soil type on qualitativedetermination of cyanogenesis for individuals of the samespecies where cyanogenic species were found on differentsubstrates all individuals tested were cyanogenic

Intra-plant variation in cyanogenic glycoside concen-tration In addition to cyanogenic polymorphism both qual-itative and quantitative within populations of cyanogenicspecies the concentration of cyanogenic glycosides variedwith leaf age and plant part In all cases where tested youngleaves or leaf tips had higher concentrations of cyanogenicglycosides than older leaves (Table 1) In some cases theconcentrations in young and old leaves differed by over anorder of magnitude (eg C sublimis and Opisthiolepisheterophylla Table 1) In terms of determining the cyano-genic phenotype of an individual this difference betweenleaves of different ages was most notable for C myrianthusand P australiana Based on FA paper tests of mature fullyexpanded leaves of these species there were frequent acy-anogenic individuals however young leaves and leaf tipssampled from the same individuals of both species routinelytested positive for cyanogenesis Furthermore amongmature trees of C myrianthus which had low levels ofcyanogenic glycosides in old leaves (6ndash8mgCNg1 d wtie considered acyanogenic) the fruit in particularhad higher concentrations of cyanogenic glycosides(gt800mgCNg1 d wt Table 1) Overall few tests weremade of seeds or other reproductive tissues as part of thesurvey however where reproductive tissues were testedfrom species with cyanogenic leaves they were typicallycyanogenic and contained higher concentrations of cyano-genic glycosides although the concentration varied withthe maturity of the fruitseed For example the immatureseed and fruit (combined) from Prunus turneriana hadgt8mgCNg1 d wt compared with mature seed alone(lt1mgCNg1 d wt) while in R javanica cyanogenic gly-coside concentrations in flesh and seed of immature fruitdecreased with maturity (Table 1 Webber and Woodrow2004) One exception was the papery wind-dispersed seedsof C sublimis which yielded negligible cyanide (Table 1)

DISCUSSION

Cyanogenic species

Despite a substantial body of early work screening Austra-lian flora including some tropical flora for cyanogenesis(eg Petrie 1912 Smith and White 1918 Finnemore andCox 1928 Hurst 1942 Webb 1948 1949) few of thespecies tested in this study had previously been testedOf the 18 species from 13 families found to be cyanogenicin this study only one F indica has previously been repor-ted as cyanogenic (Petrie 1912) Thirteen of the cyanogenicspecies are endemic to Queensland or Australia several arerestricted to small areas within north east Queensland Forexample Clerodendrum grayi is found only in a small areaon the Atherton Tableland (Miller et al 2006a) while

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1023

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R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

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Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

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(4)

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Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

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daggerSpeciesin

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whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

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siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

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aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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the field early last century reporting a number ofcyanogenic species and the specific cyanogenic consti-tuents involved (eg Petrie 1912 1913 1914 1920Finnemore and Cox 1928 1929 Finnemore andCooper 1936 1938) More than 700 species in theQueensland flora were screened by Smith and White(1918) Further phytochemical screening of the Queenslandflora was conducted by Webb (1948 1949) however therewas negligible testing among tropical taxa Importantlymany of these records are based on qualitative tests per-formed using herbarium specimens and tests of this kindusing dried material are by no means conclusive One fur-ther limitation of these data in terms of extrapolating tonatural plant communities is that negative results wereoften not reported

Tropical rainforests are of particular interest in thestudy of plant chemical defences Elevated herbivore pres-sure in tropical environments is hypothesized to havefavoured both a diverse array of defences and high levelsof investment in chemical defence (Levin 1976 Levin andYork 1978 Coley and Barone 1996 Kursar and Coley2003) Indeed in the field of plant secondary chemistry thetropical rainforest has been the focus of intense interestin co-evolutionary relationships between plants andherbivores and the extraordinary inter-specific and intra-plant variation in chemical defence strategies (Feeny1976 Levin 1976 Coley et al 1985 Coley andKursar 1996)

On the whole community-level studies of the distributionof chemical defences have focused on a small set of Asianand African rainforests (McKey et al 1978 Gartlan et al1980 Davies et al 1988 Waterman et al 1988) Inaddition these and other studies have tended to focus onC-based lsquoquantitativersquo defences (eg Coley 1983 1988)Among N-based defences the greater awareness of thefrequency of alkaloid-bearing plants in tropical and otherecosystems is a consequence of extensive phytochemicalscreening for bioactive compounds (eg Swanholm et al1960 Hartley et al 1973 Smolenski et al 1974 Collinset al 1990 Hadi and Bremmer 2001) or ecologicalresearch into specific plantndashherbivore interactions (egJanzen and Waterman 1984 Hartmann et al 1997) ratherthan systematic studies within natural communities (butsee Mali and Borges 2003)

There are some hypothesis-driven surveys for cyanogen-esis Two studies have investigated the frequency of cyano-genesis in floras investigating evolutionary and ecologicalhypotheses about exposure to herbivory (see Kaplan et al1983 Adsersen et al 1988 Adsersen and Adsersen 1993)However only the survey of Thomsen and Brimer (1997) inCosta Rican rainforest has been conducted in a systematicfashion well defined with respect to forest area and plantsize As with other N-based defences work on cyanogenictropical rainforest species worldwide is limited andthere has been no work on cyanogenesis in Australian trop-ical rainforests or any other form of chemical defence Thelack of work on cyanogenesis in diverse tropical rainforestsis further surprising as it is a readily detectable and con-stitutive chemical defence (Gleadow and Woodrow2000b)

Here we report some of the findings of a large-scalequantitative survey for cyanogenesis in Australian tropicalrainforests First we investigated the frequency of cyano-genesis and the contribution of cyanogenic species tobiomass (basal area) in lowland upland and highland trop-ical rainforest in north Queensland Australia Conductingthe survey in a standardized fashion will enable comparisonwith other communities (eg Thomsen and Brimer 1997)Secondly intra-plant and intra-population variation in con-centrations of cyanogenic glycosides was quantified Thehigh level of endemism among the Australian tropical rain-forest flora (Webb and Tracey 1981) and the number ofrainforest taxa previously untested for cyanogenesis under-score the potential for novel reports of cyanogenesis withindifferent taxonomic groups Finally therefore this studyaimed to investigate the potential taxonomic significanceof cyanogenesis reported here Overall this study aimedto expand the limited knowledge of the frequency ofcyanogenesis in the Australian flora and in natural plantcommunities

MATERIALS AND METHODS

Field sites

Field work was conducted between July 1999 andSeptember 2002 Six 200m2 plots (20 middot 10m) were estab-lished at each of five sites (total area 1200m2 per site) inlowland and upland rainforest in the tropics of north eastQueensland Australia (Fig 1) Six plots were selected fortwo reasons first the number of new species captured witheach additional plot had reached a plateau at around fivespecies and secondly it was a realistic sample size giventhe time and resources available Sites were selected tocapture maximum species diversity as on the AthertonTablelands forest type and species composition varyboth with substrate and with altitude (Tracey 1982) Fur-ther to enable comparison of forests occurring on differentsoil types two pairs of sites with similar altitude and rainfallwere selected The first pair comprised a site on soil derivedfrom basalt at Lamins Hill and one on soil derived fromgranite at Mt Nomico (Fig 1) The second pair comprisedsites on soils derived from basalt and rhyolite at LonglandsGap (Fig 1) A fifth site in lowland rainforest nearCape Tribulation and Myall Creek on soil derivedfrom metamorphic substrate was also surveyed (Fig 1)The distribution of cyanogenic species in relation toresource availability (soil nutrients) will be addressedelsewhere (see Miller 2004)

It was not possible to control strictly for logging historyall sites on the Atherton Tablelands had been selectivelylogged prior to the declaration of the Wet Tropics WorldHeritage Area in 1988 Detailed site descriptions are pro-vided in Miller (2004)

All individuals (palms trees and vines) with a diameter atbreast height (dbh) of gt5 cm were tagged and identifiedAll additional species (dbh lt5 cm) present in lower stratawith the exception of herbaceous ground species were alsotagged and identified

1018 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Samples of all cyanogenic species as well as speciesdominant at each site and some rare species were pressedand lodged at the University of Melbourne (MELU) andBrisbane (BRI) Herbaria Accession numbers whereassigned are listed (Appendix)

Climate

The climate in far north Queensland is characterizedby a marked wet season from December to April Climaterecording stations in the study area are scattered thereforedata specific to each site were not available While locatedin the tropical latitudes because of its higher altitude theclimate of the Atherton Tableland is semi-tropical Meanannual temperature within the study area on the Tablelandis 22 C (Nix 1991) with a minimum of 10 C (Hall et al1981 see Graham et al 1995) All the upland and highlandrainforests surveyed on the Atherton Tableland havehigh average annual rainfall generally in the range2000ndash3000mm plus cloud interception (Tracey 1982)For example the average annual rainfall at Lamins Hillis 3584mm based on 30 years of records from the Queens-land Bureau of Meteorology (see Osunkoya et al 1993) Incontrast the climate in the coastal lowland tropical rain-forest near Cape Tribulation is characterized by highertemperatures Mean daily temperatures range from 28 Cin January to 22 C in July and temperatures may reach

the mid to high 30 s during the summer months Averageannual rainfall is also high at 3928mm recorded at CapeTribulation (based on 65 years of records from theQueensland Bureau of Meteorology)

Sites

Upland and highland rainforest The upland rainforest atLamins Hill [172240S 1454250E altitude 850m abovesea level (asl) Fig 1] is classified as complex mesophyllvine forest on basalt (type 1b Tracey 1982 Tracey andWebb 1975) This forest type typically occurs on uplandsites (400ndash800m) on high fertility kraznozem soils derivedfrom basalt with high rainfall (Tracey 1982) It is charac-terized by a closed canopy with multiple tree layers andan uneven canopy with height ranging from 35 to 45m(Tracey 1982)

The upland rainforest at Mt Nomico (171330S1454040E 900m asl Fig 1) is within the GilliesRange State Forest and is on low nutrient soil derivedfrom granite The forest is classified as complex notophyllvine forest typical of upland granitic soils (type 6 Tracey1982 Tracey and Webb 1975)

In the highland rainforest of Longlands Gap State Forest(altitude 1100ndash1200m asl) there is a sharp boundaryin forest type defined by basalt (172770S 1452850E)and rhyolite (172730S 1452860E) parent substrates

Atherton

AthertonTableland

MalandaLonglands Gap

0 50 km

Mareeba

Cairns

Port Douglas

Daintree River

Cape TribulationMyall Creek

WesternAustralia

SouthAustralia

NorthernTerritory

Queensland

New SouthWales

Lamins HillMt Nomico

F I G 1 Location of study sites in tropical rainforest in far north east Queensland Australia There were two pairs of sites in upland rainforest on the AthertonTablelandLaminsHill (172240S 1454250E) andMtNomico (171330S 1454040E) and two sites atLonglandsGap (17270S 145280E)Afifth sitewas

in lowland rainforest near Myall Creek and Cape Tribulation (160620S 1452690E)

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1019

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On basalt the forest is complex notophyll vine forest(type 5a Tracey 1982 Tracey and Webb 1975) a foresttype characterized by an uneven canopy (20ndash40m high) andnumerous tree layers typical of basaltic cool wet uplandsand highlands (Tracey 1982) On rhyolite the forest issimple microphyll vine forest (type 9 Tracey 1982)which is common on upland granitic soils (800ndash1300m)and is characterized by an uneven canopy 20ndash25m highwith emergent Agathis atropurpurea (up to 35m)

Lowland rainforest The fifth site was in lowlandrainforest in the Daintree World Heritage Area nearCape Tribulation (Fig 1) The site was near to Thompsonand Myall Creeks (160620S 1452690E altitude 40masl) The forest is complex mesophyll vine forest (type 1aTracey 1982) the canopy is irregular from 25 to 33m inheight and supports a great diversity of species and lifeforms including many palms and lianas The floristiccomposition is patchy with considerable variation incanopy and understorey dominants over small distances(Webb et al 1972 Tracey 1982) The soil is relativelynutrient-poor red clay loam podsol derived frommetamorphic substrate

Sampling for detection of cyanogenesis

Whole leaf samples (1ndash2 g f wt) were taken from indi-viduals with a dbh gt5 cm and from all additional speciesin the lower strata until at least three individuals of eachspecies had been sampled Where possible the youngestfully expanded leaves without epiphyllous communitieswere selected however in the case of samples acquiredby pruning shears attached to extension poles or by sling-shot and rope from the canopy it was not always possibleto be selective In instances where it was not possible toobtain samples from large canopy trees samples were takenfrom nearby individuals of the same species Fresh wholeleaf samples used for cyanide testing were stored in air-tightbags on ice until tested for cyanide 2ndash6 h later using FeiglndashAnger (FA) papers (Feigl and Anger 1966) Individuals ofrare species and those with little foliage were sampled onlyonce for analysis in the laboratory where a quantitativeassay was used to test for cyanogenesis using freeze-dried ground tissue Because cyanogenesis is known tobe a polymorphic trait (eg Hughes 1991 Aikman et al1996) a minimum of three individuals of all species wastested more where species were common Owing to thediverse and heterogeneous nature of the forests multipleindividuals of each species were not always represented inthe plots Therefore for these rare species testing indivi-duals located outside the plots was required and still therewere several species for which only single samples wereobtained

A range of other factors was taken into considerationwhen sampling For example given that young leavesof tropical species in particular are typically more highlydefended than older leaves (Coley 1983 Coley and Barone1996) both young (soft expanding leaves) and old (recentfully expanded) leaves were tested for cyanogenesis wherepossible In addition depending on availability fruit andflowers from several species were tested Owing to the large

number of species and samples in the survey it was notpossible to examine seasonal trends in defence howeverindividuals of the majority of species were tested in wetand dry seasons for qualitative changes in cyanogenic sta-tus as there is some evidence for seasonal variation incyanogenesis (Seigler 1976b Janzen et al 1980 Kaplanet al 1983) Finally because edaphic factors have beenreported to affect the expression of cyanogenesis (egUrbanska 1982) species found on more than one substratetype were tested at each site

Sample collection handling and storage for quantitativeanalysis

In addition to samples for fresh cyanide tests whole leafsamples (again recent fully expanded leaves) were taken forquantification of cyanogenic glycosides All individuals ofspecies that produced a positive result and individuals ofrare species were sampled Depending on sampling condi-tions samples were either placed immediately into liquidnitrogen or placed in a sealed air-tight bag and kept on icefor 2ndash6 h until snap frozen in liquid nitrogen Frozensamples were transported to the laboratory on dry icefreeze-dried and stored on desiccant at 20 C for analysisFreeze-dried samples were ground using either a cooledIKA Labortechnic A10 Analytical Mill (Janke and KunkelStanfen Germany) or for smaller samples an Ultramat 2Dental Grinder (Southern Dental Industries Ltd BayswaterVictoria Australia)

Chemical analyses

Detection of cyanogenesis FeiglndashAnger papers Thepresence of cyanogenic compounds in fresh field sampleswas determined using FA indicator papers (Feigl andAnger 1966) FA papers were selected in preference topicrate papers because FA papers are more sensitive(Nahrstedt 1980) and less prone to giving false-positiveresults (Brinker and Seigler 1989) FA test papers wereprepared according to Brinker and Seigler (1989) BecauseFA papers can be sensitive to moisture and light (Brinkerand Seigler 1989) they were stored in the dark and ondesiccant until use

Fresh leaves (approx 1ndash2 g f wt) were crushed induplicate screw-top vials Old and young foliage sampleswere tested separately To facilitate cyanogenesis 05mL ofwater was added to one of the vials and pectinase fromRhizopus spp (Macerase Pectinase 441201 CalbiochemCalbiochem-Novabiochem Corp La Jolla CA USA)(04 g Lndash1) in 01 M TrisndashHCl (pH 68) was added to theother Pectinase has been found to have non-specific b-glyc-osidase activity (Brimer et al 1995) and therefore inthe absence of sufficient endogenous b-glycosidase enablestests for the presence of cyanogenic glycosides to be madeThe indicator papers were suspended above the tissue bymeans of the screw-top lid and vials were left at roomtemperature and checked after 12 and 24 h This 24 htime period used in other surveys (eg Dickenmann1982 Thomsen and Brimer 1997 Buhrmester et al2000 Lewis and Zona 2000) was selected to avoid spuri-ous test results due to substantial bacterial contamination

1020 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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which may occur beyond 24 h (Saupe et al 1982) Tissue ofknown cyanogenic species Prunus turneriana or Ryparosajavanica was used as a positive control Moderate tostrongly cyanogenic samples gave a positive result withina few minutes or up to a few hours while more weaklycyanogenic samples took several more hours In accordancewith the recommendations of Brinker and Seigler (1989) anew test was conducted for any samples producing a slowpositive response (24 h) in case of interference by microbialcyanogenesis In addition any samples with inconclusivecolour change were re-tested An individual was consideredcyanogenic if a positive repeatable result was obtainedand a species was considered cyanogenic if at leastone individual produced a consistent and repeatablepositive result A negative test result indicates the absenceof a cyanogenic glycoside or of the specific cyanogenicb-glycosidase or both

In the few instances where insufficient tissue was avail-able for both FA paper tests using fresh leaves and sub-sequent laboratory analysis cyanogenesis was determinedbased on the quantitative assay of freeze-dried groundleaf tissue (as described below) by comparison with anegative tissue control (eg Alstonia scholaris or Aglaiameridionalis)

In this study tests for cyanogenesis used approx1ndash2 g f wt of leaves which is larger than tissue samplestested in previous surveys (eg 50mg fwt by Lewis andZona 2000 and 200mg f wt by Thomsen and Brimer1997 Buhrmester et al 2000) According to Dickenmann(1982) who used 500mg f wt tissue a weak positive reac-tion with FA papers where part of the paper turns blueindicated approx 2ndash20mgHCNkgndash1 f wt while a strongreaction indicated gt50mgHCNkgndash1 f wt These lowervalues equate to just over 6ndash60mgHCNgndash1 d wt using aconversion based on the mean foliar water content ofseveral species in this study which was 70 In thisstudy based on fresh leaf tests and quantitative analysisof freeze-dried tissue from the same sample the thresholdsensitivity of FA papers was similar within the range5ndash8mgHCNgndash1 d wt This threshold sensitivity of FApapers corresponds well to the criteria of Adsersenet al (1988) for classifying individuals as cyanogenicwhere individuals with lt93 nmol HCNgndash1 f wt (approxim-ately equivalent to 8mgHCNgndash1 d wt) were consideredacyanogenic

Polymorphism and confirmation of acyanogenesis

For the purposes of the survey a species was consideredcyanogenic if at least one individual produced a repeatableand positive test result For a few species not all individualsproduced positive results using FA papers therefore somefurther investigation of the polymorphism for cyanogenesisin these species was conducted First to confirm the acy-anogenic character of the individuals producing negativeresults with FA papers a greater mass of freeze-dried tissue(100mg) was assayed quantitatively for the evolution ofcyanide (see below) and compared with a non-cyanogenicspecies A scholaris as a control for baseline noise in theassay Based on the criteria of Adsersen et al (1988) if

lt8mg HCNgndash1 d wt was detected then that individual wasconsidered acyanogenic

Quantification of cyanogenic glycosides

The concentration of cyanogenic glycoside in plant tis-sues was measured by trapping the cyanide (CN) liberatedfollowing hydrolysis of the glycoside in a well containing1 MNaOH (Brinker and Seigler 1989 Gleadow et al1998) Freeze-dried ground plant tissue (30ndash50mg) wasincubated for 20ndash24 h at 37 C with 1mL of 01M citratendashHCl (pH 55) Cyanide in the NaOH well was determinedusing the method of Gleadow and Woodrow (2002) adaptedfrom Brinker and Seigler (1989) for use with a photometricmicroplate reader (Labsystems Multiskan Ascent withincubator Labsystems Helsinki Finland) The method ishighly sensitive the determination of CN in NaOH isrelatively specific for cyanide and can detect as little as5mg Lndash1 (Brinker and Seigler 1989) The absorbance wasmeasured at 590 nm with NaCN as the standard

RESULTS

Survey for cyanogenesis

In total fresh leaf material from 401 woody plant speciesfrom 87 families was tested for cyanogenesis (Appendix)Cyanogenesis was found in 18 species from 13 familiesrepresenting 45 of species occurring within 1200m2 ofrainforest at each of five sites In each case the addition ofpectinase during incubation did not result in any additionalpositive results The test with added enzyme thereforeeffectively served as a replicate test for each sample Afurther two species were found to be cyanogenic butwere not included in the analysismdashthe non-woody Alocasiabrisbanensis and Helicia nortoniana which occurred out-side the plots (Table 1) The number of cyanogenic speciesat each site ranged from four to 10 The cyanogenic speciesranged in life form from climbing monocotyledons suchas the supplejack Flagellaria indica to 30m canopytrees such as the northern silky oak Cardwellia sublimisThe proportion of cyanogenic species ranged from 47 inupland rainforest at Mt Nomico to 65 in lowland rain-forest near Cape Tribulation Overall cyanogenic speciesaccounted for 73 of total basal area (range 12ndash134)The highest proportion was in highland rainforest on basaltsoil at Longlands Gap while forest on rhyolite in adjacentforest had the lowest proportion In lowland rainforestthe contribution of cyanogenic species to biomass wasalso high at 116 compared with 33 and 71 at MtNomico and Lamins Hill sites respectively

The cyanogenic species and concentrations of cyano-genic glycosides in leaves and other plant parts aresummarized in Table 1 The foliar concentrations of cyano-genic glycosides among species ranged from around8mgCNgndash1 d wtmdashthe concentration below which individu-als were considered acyanogenic based on the criteria ofAdsersen et al (1988) and the limits of FA paperdetectionmdashto very high concentrations in excess of severalmgCNgndash1 d wt (Table 1) For example fully expanded

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1021

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TABLE1

Su

mm

ary

of

the

con

cen

tra

tio

no

fcy

an

og

enic

gly

cosi

des

inle

ave

sa

nd

oth

erp

lan

tti

ssu

esfr

om

cya

no

gen

icsp

ecie

sfo

un

din

six

plo

tsin

up

lan

dh

igh

land

(U)

an

dlo

wla

nd

(L)

tro

pic

alr

ain

fore

sta

tfive

site

sh

igh

nu

trie

ntb

asa

ltsi

tes

atL

am

ins

Hil

l(B

1)

an

dL

on

gla

nd

sG

ap

(B2

)a

nd

low

nu

trie

nts

ites

on

gra

nit

ea

tMtN

om

ico

(G)

on

rhyo

lite

at

Lo

ng

lan

ds

Ga

p(R

)a

nd

on

met

am

orp

hic

sub

stra

ten

ear

Ca

pe

Tri

bu

lati

on

(M)

Concentrationofcyanogenic

glycosides

indifferentplantparts(m

gCNg1dw

t)

Form

Species

Fam

ily

Forest

Site

Leaf

Stem

Floralparts

Fruitseed

HA

loca

sia

bri

sba

nen

sis

(FM

Bailey)Domin

Araceae

UB1

ndashndash

ndashndash

TB

eils

chm

ied

iaco

llin

aBHyland

Lauraceae

UB1B2GR

Old28 4ndash1263(n

=36)

ndashndash

ndashYg1039ndash1391

TB

rom

bya

pla

tyn

emaFM

uell

Rutaceae

LM

156 4ndash1285(n

=20)

ndashndash

ndash0ndash10 5

(n=27)

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

UL

B1B1GRM

Old11 5ndash69 8

(n=21)

ndashBud779ndash851

Seedapprox8

Yg733 1tips219 1

Flwr130ndash334

TC

leis

tanth

us

myr

ian

thu

s(H

assk)Kurz

Euphorbiaceae

LM

Old1 1ndash17 3

(n=41)

ndashFlwr30 7

Mature160ndash377

Yg78 4ndash80 4

(n=3)

Immature821

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

UB1B2G

Old1325ndash4800(n

=6)

ndashBud733

ndashFlwr440ndash942

TE

laeo

carp

us

seri

cop

etalu

sFM

uell

Elaeocarpaceae

UGR

1100ndash5056(n

=10)

Sdlg1739ndash2679

ndash1028

Tree

135

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

UB1B2

10ndash81(n

=3)

ndashndash

ndashV

Fla

gel

lari

ain

dic

aL

Flagellariaceae

UL

B1GM

11ndash177 3

(n=6)

ndashndash

ndashT

Hel

icia

aust

rala

sica

FM

uell

Proteaceae

LM

42 2ndash155 2

(n=8)dagger

ndashndash

ndashT

Hel

icia

bla

keiForeman

Proteaceae

UB1

Mean17 9

(n=2)

ndashndash

ndashT

Hel

icia

no

rto

nia

na

(FM

Bailey)FM

Bailey

Proteaceae

LM

ndashndash

ndashndash

VP

ass

iflo

rasp(K

urandaBH12896)

Passifloraceae

LM

313ndash922(n

=4)

ndashndash

ndashT

Mis

cho

carp

us

exa

ng

ula

tus

(FM

uell)Radlk

Sapindaceae

UB1

Old133yg222(n

=1)

ndashndash

ndash

TM

isch

oca

rpu

sg

ran

dis

sim

us

(FM

uell)Radlk

Sapindaceae

UL

GM

49ndash680(n

=2)

761ndash2006(n

=4)z

ndashndash

ndash

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

UB1B2

Old10ndash208(n

=7)

ndashndash

ndashYg2000

VP

ars

on

sia

lati

foli

a(Benth)

STBlake

Apocynaceae

UB1GR

765ndash4835(n

=3)

ndashndash

ndash

TP

oly

scia

sa

ust

rali

an

a(FM

uell)Philipson

Araliaceae

UL

B1B2GRM

Oldlt6

8(n

=30)10ndash28 5

(n=16)

ndashndash

ndashYg10 8ndash29 6tips259

TP

runu

stu

rner

ian

a(FM

Bailey)Kalkman

Rosaceae

UL

B1M

old2472ndash4888(n

=4)

560ndash1100(n

=2)

ndashMature

seed400

Yg3128ndash6464Tip6000ndash8498

Rootapprox2000

Ygfruitseed8950

TR

ypa

rosa

java

nic

a(Blume)

Kurz

exKoordamp

Valeton

Achariaceae

LM

1749(n

=2)

ndashndash

Mature

seed5430

Yglf2560(n

=5)

Ygseed7760(n

=5)

Meansandranges

ofcyanogenicglycosides

concentrationaregiven

forfullyexpanded

(old)leaveswherenotspecifiedandforsomeyoung(yg)leavesfor

nindividualsLifeform

herb(H

)tree

(T)vine(V

)Cyanogenic

glycosideconcentrationsweremeasuredas

evolved

cyanidefollowinghydrolysisoffreeze-dried

tissuewithouttheadditionofnon-specificb-glycosidase(egem

ulsinpectinase)

A

loca

sia

bri

sba

nen

sis(a

herb)and

Hel

icia

nort

onia

nawhichoccurred

outsidetheplotswerenotincluded

intheanalysisnorsampledforquantitativeanalysis

Concentrationsofcyanogenic

glycosides

inseedflower

andyoungleaves

of

Ryp

aro

saja

van

icadetermined

byBLWebber

(unpubldata2004Webber

andWoodrow2004)

daggerOnly

oneindividual

of

Hel

icia

aust

rala

sica

occurred

within

theplotsseven

additional

saplingstreeswereopportunisticallytested

externalto

theplotsallwerecyanogenic

zOnly

oneindividual

of

Mis

cho

carp

us

gra

nd

issi

mu

soccurred

insideplotsat

MtNomicoadditional

individualsweresampledoutsideplotsincludingfourin

lowlandrainforestallwerecyanogenic

1022 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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leaves of numerous individuals of Polysciasaustraliana tested negative for cyanide and containedlt8mgCNgndash1 d wt while others tested positive and werein the range of 10ndash28 mgCNgndash1 d wt Even in the weaklycyanogenic samples only in very few cases did the colora-tion of the test paper change from 12 to 24 h in a qualit-ative sense Individuals of numerous species had highconcentrations of cyanogenic glycosides in excess of1mgCNgndash1 d wt (eg Mischocarpus grandissimusBrombya platynema and Beilschmiedia collina) and severalspecies contained extremely high concentrations of cyano-genic glycosides in mature tree leaves Notably fullyexpanded leaves from the tree species Elaeocarpus serico-petalus Clerodendrum grayi and Prunus turneriana hadconcentrations of cyanogenic glycosides up to 52 49and 48mgCNgndash1 d wt respectively (Table 1)

Qualitative and quantitative variation in cyanogenesis

Intra-population variation in cyanogenic glycosides Forthe majority of species initially identified as being cyano-genic all individuals tested were cyanogenic however theconcentrations of cyanogenic glycosides varied markedlybetween conspecific individuals (Table 1) For example allsaplings and trees of B collina at each site tested positivefor cyanogenesis but the concentration of cyanogenic glyc-osides in fully expanded leaves from saplings (n = 19)ranged from 232 to 12634mgCNgndash1 d wt at Mt Nomicoalone where B collina was most abundant In contrast tothis quantitative phenotypic variation in cyanogenesis con-specific individuals of a small number of species producedboth positive and negative FA paper test results Thisapparent qualitative polymorphism was most notable inB platynema a subcanopy tree species restricted to thelowland rainforest Within the population of B platynemaapprox 50 of individuals tested produced negativeFA test paper results for both young and old leaves andhad cyanogenic glycoside concentrations in the range06ndash68mgCNgndash1 d wt (Table 1) The addition ofbuffered pectinase during testing did not alter the qualitativeFA paper test result for any individual Among cyanogenicB platynema the concentrations of cyanogenic glyco-sides varied over orders of magnitude from 105 to12859mgCN gndash1 d wt (Table 1)

Negative results with FA papers were obtained for twoother species Cleistanthus myrianthus and Polysciasaustraliana Qualitative and quantitative analysis of oldleaves from individuals of these species indicated a highfrequency of acyanogenic individuals however unlikeB platynema the phenotypic expression of cyanogenesisvaried qualitatively with leaf age such that young leavesand leaf tips from these same individuals were consistentlycyanogenic aswere reproductive tissues fromC myrianthus(Table 1)

Even among less common species with smallsample sizes the concentrations of cyanogenic glycosidesvaried markedly between individuals For example inMischocarpus grandissimus the concentration amongsix individuals from two sites ranged from 49 to680mgCNgndash1 d wt at Mt Nomico and from 637 to

2006mgCNg1 d wt in lowland rainforest and amongthree individuals of the woody vine Parsonsia latifoliaconcentrations ranged from 765 to 4835 mgCNg1 d wtThere was no detectable affect of soil type on qualitativedetermination of cyanogenesis for individuals of the samespecies where cyanogenic species were found on differentsubstrates all individuals tested were cyanogenic

Intra-plant variation in cyanogenic glycoside concen-tration In addition to cyanogenic polymorphism both qual-itative and quantitative within populations of cyanogenicspecies the concentration of cyanogenic glycosides variedwith leaf age and plant part In all cases where tested youngleaves or leaf tips had higher concentrations of cyanogenicglycosides than older leaves (Table 1) In some cases theconcentrations in young and old leaves differed by over anorder of magnitude (eg C sublimis and Opisthiolepisheterophylla Table 1) In terms of determining the cyano-genic phenotype of an individual this difference betweenleaves of different ages was most notable for C myrianthusand P australiana Based on FA paper tests of mature fullyexpanded leaves of these species there were frequent acy-anogenic individuals however young leaves and leaf tipssampled from the same individuals of both species routinelytested positive for cyanogenesis Furthermore amongmature trees of C myrianthus which had low levels ofcyanogenic glycosides in old leaves (6ndash8mgCNg1 d wtie considered acyanogenic) the fruit in particularhad higher concentrations of cyanogenic glycosides(gt800mgCNg1 d wt Table 1) Overall few tests weremade of seeds or other reproductive tissues as part of thesurvey however where reproductive tissues were testedfrom species with cyanogenic leaves they were typicallycyanogenic and contained higher concentrations of cyano-genic glycosides although the concentration varied withthe maturity of the fruitseed For example the immatureseed and fruit (combined) from Prunus turneriana hadgt8mgCNg1 d wt compared with mature seed alone(lt1mgCNg1 d wt) while in R javanica cyanogenic gly-coside concentrations in flesh and seed of immature fruitdecreased with maturity (Table 1 Webber and Woodrow2004) One exception was the papery wind-dispersed seedsof C sublimis which yielded negligible cyanide (Table 1)

DISCUSSION

Cyanogenic species

Despite a substantial body of early work screening Austra-lian flora including some tropical flora for cyanogenesis(eg Petrie 1912 Smith and White 1918 Finnemore andCox 1928 Hurst 1942 Webb 1948 1949) few of thespecies tested in this study had previously been testedOf the 18 species from 13 families found to be cyanogenicin this study only one F indica has previously been repor-ted as cyanogenic (Petrie 1912) Thirteen of the cyanogenicspecies are endemic to Queensland or Australia several arerestricted to small areas within north east Queensland Forexample Clerodendrum grayi is found only in a small areaon the Atherton Tableland (Miller et al 2006a) while

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1023

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R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Samples of all cyanogenic species as well as speciesdominant at each site and some rare species were pressedand lodged at the University of Melbourne (MELU) andBrisbane (BRI) Herbaria Accession numbers whereassigned are listed (Appendix)

Climate

The climate in far north Queensland is characterizedby a marked wet season from December to April Climaterecording stations in the study area are scattered thereforedata specific to each site were not available While locatedin the tropical latitudes because of its higher altitude theclimate of the Atherton Tableland is semi-tropical Meanannual temperature within the study area on the Tablelandis 22 C (Nix 1991) with a minimum of 10 C (Hall et al1981 see Graham et al 1995) All the upland and highlandrainforests surveyed on the Atherton Tableland havehigh average annual rainfall generally in the range2000ndash3000mm plus cloud interception (Tracey 1982)For example the average annual rainfall at Lamins Hillis 3584mm based on 30 years of records from the Queens-land Bureau of Meteorology (see Osunkoya et al 1993) Incontrast the climate in the coastal lowland tropical rain-forest near Cape Tribulation is characterized by highertemperatures Mean daily temperatures range from 28 Cin January to 22 C in July and temperatures may reach

the mid to high 30 s during the summer months Averageannual rainfall is also high at 3928mm recorded at CapeTribulation (based on 65 years of records from theQueensland Bureau of Meteorology)

Sites

Upland and highland rainforest The upland rainforest atLamins Hill [172240S 1454250E altitude 850m abovesea level (asl) Fig 1] is classified as complex mesophyllvine forest on basalt (type 1b Tracey 1982 Tracey andWebb 1975) This forest type typically occurs on uplandsites (400ndash800m) on high fertility kraznozem soils derivedfrom basalt with high rainfall (Tracey 1982) It is charac-terized by a closed canopy with multiple tree layers andan uneven canopy with height ranging from 35 to 45m(Tracey 1982)

The upland rainforest at Mt Nomico (171330S1454040E 900m asl Fig 1) is within the GilliesRange State Forest and is on low nutrient soil derivedfrom granite The forest is classified as complex notophyllvine forest typical of upland granitic soils (type 6 Tracey1982 Tracey and Webb 1975)

In the highland rainforest of Longlands Gap State Forest(altitude 1100ndash1200m asl) there is a sharp boundaryin forest type defined by basalt (172770S 1452850E)and rhyolite (172730S 1452860E) parent substrates

Atherton

AthertonTableland

MalandaLonglands Gap

0 50 km

Mareeba

Cairns

Port Douglas

Daintree River

Cape TribulationMyall Creek

WesternAustralia

SouthAustralia

NorthernTerritory

Queensland

New SouthWales

Lamins HillMt Nomico

F I G 1 Location of study sites in tropical rainforest in far north east Queensland Australia There were two pairs of sites in upland rainforest on the AthertonTablelandLaminsHill (172240S 1454250E) andMtNomico (171330S 1454040E) and two sites atLonglandsGap (17270S 145280E)Afifth sitewas

in lowland rainforest near Myall Creek and Cape Tribulation (160620S 1452690E)

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1019

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On basalt the forest is complex notophyll vine forest(type 5a Tracey 1982 Tracey and Webb 1975) a foresttype characterized by an uneven canopy (20ndash40m high) andnumerous tree layers typical of basaltic cool wet uplandsand highlands (Tracey 1982) On rhyolite the forest issimple microphyll vine forest (type 9 Tracey 1982)which is common on upland granitic soils (800ndash1300m)and is characterized by an uneven canopy 20ndash25m highwith emergent Agathis atropurpurea (up to 35m)

Lowland rainforest The fifth site was in lowlandrainforest in the Daintree World Heritage Area nearCape Tribulation (Fig 1) The site was near to Thompsonand Myall Creeks (160620S 1452690E altitude 40masl) The forest is complex mesophyll vine forest (type 1aTracey 1982) the canopy is irregular from 25 to 33m inheight and supports a great diversity of species and lifeforms including many palms and lianas The floristiccomposition is patchy with considerable variation incanopy and understorey dominants over small distances(Webb et al 1972 Tracey 1982) The soil is relativelynutrient-poor red clay loam podsol derived frommetamorphic substrate

Sampling for detection of cyanogenesis

Whole leaf samples (1ndash2 g f wt) were taken from indi-viduals with a dbh gt5 cm and from all additional speciesin the lower strata until at least three individuals of eachspecies had been sampled Where possible the youngestfully expanded leaves without epiphyllous communitieswere selected however in the case of samples acquiredby pruning shears attached to extension poles or by sling-shot and rope from the canopy it was not always possibleto be selective In instances where it was not possible toobtain samples from large canopy trees samples were takenfrom nearby individuals of the same species Fresh wholeleaf samples used for cyanide testing were stored in air-tightbags on ice until tested for cyanide 2ndash6 h later using FeiglndashAnger (FA) papers (Feigl and Anger 1966) Individuals ofrare species and those with little foliage were sampled onlyonce for analysis in the laboratory where a quantitativeassay was used to test for cyanogenesis using freeze-dried ground tissue Because cyanogenesis is known tobe a polymorphic trait (eg Hughes 1991 Aikman et al1996) a minimum of three individuals of all species wastested more where species were common Owing to thediverse and heterogeneous nature of the forests multipleindividuals of each species were not always represented inthe plots Therefore for these rare species testing indivi-duals located outside the plots was required and still therewere several species for which only single samples wereobtained

A range of other factors was taken into considerationwhen sampling For example given that young leavesof tropical species in particular are typically more highlydefended than older leaves (Coley 1983 Coley and Barone1996) both young (soft expanding leaves) and old (recentfully expanded) leaves were tested for cyanogenesis wherepossible In addition depending on availability fruit andflowers from several species were tested Owing to the large

number of species and samples in the survey it was notpossible to examine seasonal trends in defence howeverindividuals of the majority of species were tested in wetand dry seasons for qualitative changes in cyanogenic sta-tus as there is some evidence for seasonal variation incyanogenesis (Seigler 1976b Janzen et al 1980 Kaplanet al 1983) Finally because edaphic factors have beenreported to affect the expression of cyanogenesis (egUrbanska 1982) species found on more than one substratetype were tested at each site

Sample collection handling and storage for quantitativeanalysis

In addition to samples for fresh cyanide tests whole leafsamples (again recent fully expanded leaves) were taken forquantification of cyanogenic glycosides All individuals ofspecies that produced a positive result and individuals ofrare species were sampled Depending on sampling condi-tions samples were either placed immediately into liquidnitrogen or placed in a sealed air-tight bag and kept on icefor 2ndash6 h until snap frozen in liquid nitrogen Frozensamples were transported to the laboratory on dry icefreeze-dried and stored on desiccant at 20 C for analysisFreeze-dried samples were ground using either a cooledIKA Labortechnic A10 Analytical Mill (Janke and KunkelStanfen Germany) or for smaller samples an Ultramat 2Dental Grinder (Southern Dental Industries Ltd BayswaterVictoria Australia)

Chemical analyses

Detection of cyanogenesis FeiglndashAnger papers Thepresence of cyanogenic compounds in fresh field sampleswas determined using FA indicator papers (Feigl andAnger 1966) FA papers were selected in preference topicrate papers because FA papers are more sensitive(Nahrstedt 1980) and less prone to giving false-positiveresults (Brinker and Seigler 1989) FA test papers wereprepared according to Brinker and Seigler (1989) BecauseFA papers can be sensitive to moisture and light (Brinkerand Seigler 1989) they were stored in the dark and ondesiccant until use

Fresh leaves (approx 1ndash2 g f wt) were crushed induplicate screw-top vials Old and young foliage sampleswere tested separately To facilitate cyanogenesis 05mL ofwater was added to one of the vials and pectinase fromRhizopus spp (Macerase Pectinase 441201 CalbiochemCalbiochem-Novabiochem Corp La Jolla CA USA)(04 g Lndash1) in 01 M TrisndashHCl (pH 68) was added to theother Pectinase has been found to have non-specific b-glyc-osidase activity (Brimer et al 1995) and therefore inthe absence of sufficient endogenous b-glycosidase enablestests for the presence of cyanogenic glycosides to be madeThe indicator papers were suspended above the tissue bymeans of the screw-top lid and vials were left at roomtemperature and checked after 12 and 24 h This 24 htime period used in other surveys (eg Dickenmann1982 Thomsen and Brimer 1997 Buhrmester et al2000 Lewis and Zona 2000) was selected to avoid spuri-ous test results due to substantial bacterial contamination

1020 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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which may occur beyond 24 h (Saupe et al 1982) Tissue ofknown cyanogenic species Prunus turneriana or Ryparosajavanica was used as a positive control Moderate tostrongly cyanogenic samples gave a positive result withina few minutes or up to a few hours while more weaklycyanogenic samples took several more hours In accordancewith the recommendations of Brinker and Seigler (1989) anew test was conducted for any samples producing a slowpositive response (24 h) in case of interference by microbialcyanogenesis In addition any samples with inconclusivecolour change were re-tested An individual was consideredcyanogenic if a positive repeatable result was obtainedand a species was considered cyanogenic if at leastone individual produced a consistent and repeatablepositive result A negative test result indicates the absenceof a cyanogenic glycoside or of the specific cyanogenicb-glycosidase or both

In the few instances where insufficient tissue was avail-able for both FA paper tests using fresh leaves and sub-sequent laboratory analysis cyanogenesis was determinedbased on the quantitative assay of freeze-dried groundleaf tissue (as described below) by comparison with anegative tissue control (eg Alstonia scholaris or Aglaiameridionalis)

In this study tests for cyanogenesis used approx1ndash2 g f wt of leaves which is larger than tissue samplestested in previous surveys (eg 50mg fwt by Lewis andZona 2000 and 200mg f wt by Thomsen and Brimer1997 Buhrmester et al 2000) According to Dickenmann(1982) who used 500mg f wt tissue a weak positive reac-tion with FA papers where part of the paper turns blueindicated approx 2ndash20mgHCNkgndash1 f wt while a strongreaction indicated gt50mgHCNkgndash1 f wt These lowervalues equate to just over 6ndash60mgHCNgndash1 d wt using aconversion based on the mean foliar water content ofseveral species in this study which was 70 In thisstudy based on fresh leaf tests and quantitative analysisof freeze-dried tissue from the same sample the thresholdsensitivity of FA papers was similar within the range5ndash8mgHCNgndash1 d wt This threshold sensitivity of FApapers corresponds well to the criteria of Adsersenet al (1988) for classifying individuals as cyanogenicwhere individuals with lt93 nmol HCNgndash1 f wt (approxim-ately equivalent to 8mgHCNgndash1 d wt) were consideredacyanogenic

Polymorphism and confirmation of acyanogenesis

For the purposes of the survey a species was consideredcyanogenic if at least one individual produced a repeatableand positive test result For a few species not all individualsproduced positive results using FA papers therefore somefurther investigation of the polymorphism for cyanogenesisin these species was conducted First to confirm the acy-anogenic character of the individuals producing negativeresults with FA papers a greater mass of freeze-dried tissue(100mg) was assayed quantitatively for the evolution ofcyanide (see below) and compared with a non-cyanogenicspecies A scholaris as a control for baseline noise in theassay Based on the criteria of Adsersen et al (1988) if

lt8mg HCNgndash1 d wt was detected then that individual wasconsidered acyanogenic

Quantification of cyanogenic glycosides

The concentration of cyanogenic glycoside in plant tis-sues was measured by trapping the cyanide (CN) liberatedfollowing hydrolysis of the glycoside in a well containing1 MNaOH (Brinker and Seigler 1989 Gleadow et al1998) Freeze-dried ground plant tissue (30ndash50mg) wasincubated for 20ndash24 h at 37 C with 1mL of 01M citratendashHCl (pH 55) Cyanide in the NaOH well was determinedusing the method of Gleadow and Woodrow (2002) adaptedfrom Brinker and Seigler (1989) for use with a photometricmicroplate reader (Labsystems Multiskan Ascent withincubator Labsystems Helsinki Finland) The method ishighly sensitive the determination of CN in NaOH isrelatively specific for cyanide and can detect as little as5mg Lndash1 (Brinker and Seigler 1989) The absorbance wasmeasured at 590 nm with NaCN as the standard

RESULTS

Survey for cyanogenesis

In total fresh leaf material from 401 woody plant speciesfrom 87 families was tested for cyanogenesis (Appendix)Cyanogenesis was found in 18 species from 13 familiesrepresenting 45 of species occurring within 1200m2 ofrainforest at each of five sites In each case the addition ofpectinase during incubation did not result in any additionalpositive results The test with added enzyme thereforeeffectively served as a replicate test for each sample Afurther two species were found to be cyanogenic butwere not included in the analysismdashthe non-woody Alocasiabrisbanensis and Helicia nortoniana which occurred out-side the plots (Table 1) The number of cyanogenic speciesat each site ranged from four to 10 The cyanogenic speciesranged in life form from climbing monocotyledons suchas the supplejack Flagellaria indica to 30m canopytrees such as the northern silky oak Cardwellia sublimisThe proportion of cyanogenic species ranged from 47 inupland rainforest at Mt Nomico to 65 in lowland rain-forest near Cape Tribulation Overall cyanogenic speciesaccounted for 73 of total basal area (range 12ndash134)The highest proportion was in highland rainforest on basaltsoil at Longlands Gap while forest on rhyolite in adjacentforest had the lowest proportion In lowland rainforestthe contribution of cyanogenic species to biomass wasalso high at 116 compared with 33 and 71 at MtNomico and Lamins Hill sites respectively

The cyanogenic species and concentrations of cyano-genic glycosides in leaves and other plant parts aresummarized in Table 1 The foliar concentrations of cyano-genic glycosides among species ranged from around8mgCNgndash1 d wtmdashthe concentration below which individu-als were considered acyanogenic based on the criteria ofAdsersen et al (1988) and the limits of FA paperdetectionmdashto very high concentrations in excess of severalmgCNgndash1 d wt (Table 1) For example fully expanded

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1021

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TABLE1

Su

mm

ary

of

the

con

cen

tra

tio

no

fcy

an

og

enic

gly

cosi

des

inle

ave

sa

nd

oth

erp

lan

tti

ssu

esfr

om

cya

no

gen

icsp

ecie

sfo

un

din

six

plo

tsin

up

lan

dh

igh

land

(U)

an

dlo

wla

nd

(L)

tro

pic

alr

ain

fore

sta

tfive

site

sh

igh

nu

trie

ntb

asa

ltsi

tes

atL

am

ins

Hil

l(B

1)

an

dL

on

gla

nd

sG

ap

(B2

)a

nd

low

nu

trie

nts

ites

on

gra

nit

ea

tMtN

om

ico

(G)

on

rhyo

lite

at

Lo

ng

lan

ds

Ga

p(R

)a

nd

on

met

am

orp

hic

sub

stra

ten

ear

Ca

pe

Tri

bu

lati

on

(M)

Concentrationofcyanogenic

glycosides

indifferentplantparts(m

gCNg1dw

t)

Form

Species

Fam

ily

Forest

Site

Leaf

Stem

Floralparts

Fruitseed

HA

loca

sia

bri

sba

nen

sis

(FM

Bailey)Domin

Araceae

UB1

ndashndash

ndashndash

TB

eils

chm

ied

iaco

llin

aBHyland

Lauraceae

UB1B2GR

Old28 4ndash1263(n

=36)

ndashndash

ndashYg1039ndash1391

TB

rom

bya

pla

tyn

emaFM

uell

Rutaceae

LM

156 4ndash1285(n

=20)

ndashndash

ndash0ndash10 5

(n=27)

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

UL

B1B1GRM

Old11 5ndash69 8

(n=21)

ndashBud779ndash851

Seedapprox8

Yg733 1tips219 1

Flwr130ndash334

TC

leis

tanth

us

myr

ian

thu

s(H

assk)Kurz

Euphorbiaceae

LM

Old1 1ndash17 3

(n=41)

ndashFlwr30 7

Mature160ndash377

Yg78 4ndash80 4

(n=3)

Immature821

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

UB1B2G

Old1325ndash4800(n

=6)

ndashBud733

ndashFlwr440ndash942

TE

laeo

carp

us

seri

cop

etalu

sFM

uell

Elaeocarpaceae

UGR

1100ndash5056(n

=10)

Sdlg1739ndash2679

ndash1028

Tree

135

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

UB1B2

10ndash81(n

=3)

ndashndash

ndashV

Fla

gel

lari

ain

dic

aL

Flagellariaceae

UL

B1GM

11ndash177 3

(n=6)

ndashndash

ndashT

Hel

icia

aust

rala

sica

FM

uell

Proteaceae

LM

42 2ndash155 2

(n=8)dagger

ndashndash

ndashT

Hel

icia

bla

keiForeman

Proteaceae

UB1

Mean17 9

(n=2)

ndashndash

ndashT

Hel

icia

no

rto

nia

na

(FM

Bailey)FM

Bailey

Proteaceae

LM

ndashndash

ndashndash

VP

ass

iflo

rasp(K

urandaBH12896)

Passifloraceae

LM

313ndash922(n

=4)

ndashndash

ndashT

Mis

cho

carp

us

exa

ng

ula

tus

(FM

uell)Radlk

Sapindaceae

UB1

Old133yg222(n

=1)

ndashndash

ndash

TM

isch

oca

rpu

sg

ran

dis

sim

us

(FM

uell)Radlk

Sapindaceae

UL

GM

49ndash680(n

=2)

761ndash2006(n

=4)z

ndashndash

ndash

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

UB1B2

Old10ndash208(n

=7)

ndashndash

ndashYg2000

VP

ars

on

sia

lati

foli

a(Benth)

STBlake

Apocynaceae

UB1GR

765ndash4835(n

=3)

ndashndash

ndash

TP

oly

scia

sa

ust

rali

an

a(FM

uell)Philipson

Araliaceae

UL

B1B2GRM

Oldlt6

8(n

=30)10ndash28 5

(n=16)

ndashndash

ndashYg10 8ndash29 6tips259

TP

runu

stu

rner

ian

a(FM

Bailey)Kalkman

Rosaceae

UL

B1M

old2472ndash4888(n

=4)

560ndash1100(n

=2)

ndashMature

seed400

Yg3128ndash6464Tip6000ndash8498

Rootapprox2000

Ygfruitseed8950

TR

ypa

rosa

java

nic

a(Blume)

Kurz

exKoordamp

Valeton

Achariaceae

LM

1749(n

=2)

ndashndash

Mature

seed5430

Yglf2560(n

=5)

Ygseed7760(n

=5)

Meansandranges

ofcyanogenicglycosides

concentrationaregiven

forfullyexpanded

(old)leaveswherenotspecifiedandforsomeyoung(yg)leavesfor

nindividualsLifeform

herb(H

)tree

(T)vine(V

)Cyanogenic

glycosideconcentrationsweremeasuredas

evolved

cyanidefollowinghydrolysisoffreeze-dried

tissuewithouttheadditionofnon-specificb-glycosidase(egem

ulsinpectinase)

A

loca

sia

bri

sba

nen

sis(a

herb)and

Hel

icia

nort

onia

nawhichoccurred

outsidetheplotswerenotincluded

intheanalysisnorsampledforquantitativeanalysis

Concentrationsofcyanogenic

glycosides

inseedflower

andyoungleaves

of

Ryp

aro

saja

van

icadetermined

byBLWebber

(unpubldata2004Webber

andWoodrow2004)

daggerOnly

oneindividual

of

Hel

icia

aust

rala

sica

occurred

within

theplotsseven

additional

saplingstreeswereopportunisticallytested

externalto

theplotsallwerecyanogenic

zOnly

oneindividual

of

Mis

cho

carp

us

gra

nd

issi

mu

soccurred

insideplotsat

MtNomicoadditional

individualsweresampledoutsideplotsincludingfourin

lowlandrainforestallwerecyanogenic

1022 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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leaves of numerous individuals of Polysciasaustraliana tested negative for cyanide and containedlt8mgCNgndash1 d wt while others tested positive and werein the range of 10ndash28 mgCNgndash1 d wt Even in the weaklycyanogenic samples only in very few cases did the colora-tion of the test paper change from 12 to 24 h in a qualit-ative sense Individuals of numerous species had highconcentrations of cyanogenic glycosides in excess of1mgCNgndash1 d wt (eg Mischocarpus grandissimusBrombya platynema and Beilschmiedia collina) and severalspecies contained extremely high concentrations of cyano-genic glycosides in mature tree leaves Notably fullyexpanded leaves from the tree species Elaeocarpus serico-petalus Clerodendrum grayi and Prunus turneriana hadconcentrations of cyanogenic glycosides up to 52 49and 48mgCNgndash1 d wt respectively (Table 1)

Qualitative and quantitative variation in cyanogenesis

Intra-population variation in cyanogenic glycosides Forthe majority of species initially identified as being cyano-genic all individuals tested were cyanogenic however theconcentrations of cyanogenic glycosides varied markedlybetween conspecific individuals (Table 1) For example allsaplings and trees of B collina at each site tested positivefor cyanogenesis but the concentration of cyanogenic glyc-osides in fully expanded leaves from saplings (n = 19)ranged from 232 to 12634mgCNgndash1 d wt at Mt Nomicoalone where B collina was most abundant In contrast tothis quantitative phenotypic variation in cyanogenesis con-specific individuals of a small number of species producedboth positive and negative FA paper test results Thisapparent qualitative polymorphism was most notable inB platynema a subcanopy tree species restricted to thelowland rainforest Within the population of B platynemaapprox 50 of individuals tested produced negativeFA test paper results for both young and old leaves andhad cyanogenic glycoside concentrations in the range06ndash68mgCNgndash1 d wt (Table 1) The addition ofbuffered pectinase during testing did not alter the qualitativeFA paper test result for any individual Among cyanogenicB platynema the concentrations of cyanogenic glyco-sides varied over orders of magnitude from 105 to12859mgCN gndash1 d wt (Table 1)

Negative results with FA papers were obtained for twoother species Cleistanthus myrianthus and Polysciasaustraliana Qualitative and quantitative analysis of oldleaves from individuals of these species indicated a highfrequency of acyanogenic individuals however unlikeB platynema the phenotypic expression of cyanogenesisvaried qualitatively with leaf age such that young leavesand leaf tips from these same individuals were consistentlycyanogenic aswere reproductive tissues fromC myrianthus(Table 1)

Even among less common species with smallsample sizes the concentrations of cyanogenic glycosidesvaried markedly between individuals For example inMischocarpus grandissimus the concentration amongsix individuals from two sites ranged from 49 to680mgCNgndash1 d wt at Mt Nomico and from 637 to

2006mgCNg1 d wt in lowland rainforest and amongthree individuals of the woody vine Parsonsia latifoliaconcentrations ranged from 765 to 4835 mgCNg1 d wtThere was no detectable affect of soil type on qualitativedetermination of cyanogenesis for individuals of the samespecies where cyanogenic species were found on differentsubstrates all individuals tested were cyanogenic

Intra-plant variation in cyanogenic glycoside concen-tration In addition to cyanogenic polymorphism both qual-itative and quantitative within populations of cyanogenicspecies the concentration of cyanogenic glycosides variedwith leaf age and plant part In all cases where tested youngleaves or leaf tips had higher concentrations of cyanogenicglycosides than older leaves (Table 1) In some cases theconcentrations in young and old leaves differed by over anorder of magnitude (eg C sublimis and Opisthiolepisheterophylla Table 1) In terms of determining the cyano-genic phenotype of an individual this difference betweenleaves of different ages was most notable for C myrianthusand P australiana Based on FA paper tests of mature fullyexpanded leaves of these species there were frequent acy-anogenic individuals however young leaves and leaf tipssampled from the same individuals of both species routinelytested positive for cyanogenesis Furthermore amongmature trees of C myrianthus which had low levels ofcyanogenic glycosides in old leaves (6ndash8mgCNg1 d wtie considered acyanogenic) the fruit in particularhad higher concentrations of cyanogenic glycosides(gt800mgCNg1 d wt Table 1) Overall few tests weremade of seeds or other reproductive tissues as part of thesurvey however where reproductive tissues were testedfrom species with cyanogenic leaves they were typicallycyanogenic and contained higher concentrations of cyano-genic glycosides although the concentration varied withthe maturity of the fruitseed For example the immatureseed and fruit (combined) from Prunus turneriana hadgt8mgCNg1 d wt compared with mature seed alone(lt1mgCNg1 d wt) while in R javanica cyanogenic gly-coside concentrations in flesh and seed of immature fruitdecreased with maturity (Table 1 Webber and Woodrow2004) One exception was the papery wind-dispersed seedsof C sublimis which yielded negligible cyanide (Table 1)

DISCUSSION

Cyanogenic species

Despite a substantial body of early work screening Austra-lian flora including some tropical flora for cyanogenesis(eg Petrie 1912 Smith and White 1918 Finnemore andCox 1928 Hurst 1942 Webb 1948 1949) few of thespecies tested in this study had previously been testedOf the 18 species from 13 families found to be cyanogenicin this study only one F indica has previously been repor-ted as cyanogenic (Petrie 1912) Thirteen of the cyanogenicspecies are endemic to Queensland or Australia several arerestricted to small areas within north east Queensland Forexample Clerodendrum grayi is found only in a small areaon the Atherton Tableland (Miller et al 2006a) while

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1023

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R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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On basalt the forest is complex notophyll vine forest(type 5a Tracey 1982 Tracey and Webb 1975) a foresttype characterized by an uneven canopy (20ndash40m high) andnumerous tree layers typical of basaltic cool wet uplandsand highlands (Tracey 1982) On rhyolite the forest issimple microphyll vine forest (type 9 Tracey 1982)which is common on upland granitic soils (800ndash1300m)and is characterized by an uneven canopy 20ndash25m highwith emergent Agathis atropurpurea (up to 35m)

Lowland rainforest The fifth site was in lowlandrainforest in the Daintree World Heritage Area nearCape Tribulation (Fig 1) The site was near to Thompsonand Myall Creeks (160620S 1452690E altitude 40masl) The forest is complex mesophyll vine forest (type 1aTracey 1982) the canopy is irregular from 25 to 33m inheight and supports a great diversity of species and lifeforms including many palms and lianas The floristiccomposition is patchy with considerable variation incanopy and understorey dominants over small distances(Webb et al 1972 Tracey 1982) The soil is relativelynutrient-poor red clay loam podsol derived frommetamorphic substrate

Sampling for detection of cyanogenesis

Whole leaf samples (1ndash2 g f wt) were taken from indi-viduals with a dbh gt5 cm and from all additional speciesin the lower strata until at least three individuals of eachspecies had been sampled Where possible the youngestfully expanded leaves without epiphyllous communitieswere selected however in the case of samples acquiredby pruning shears attached to extension poles or by sling-shot and rope from the canopy it was not always possibleto be selective In instances where it was not possible toobtain samples from large canopy trees samples were takenfrom nearby individuals of the same species Fresh wholeleaf samples used for cyanide testing were stored in air-tightbags on ice until tested for cyanide 2ndash6 h later using FeiglndashAnger (FA) papers (Feigl and Anger 1966) Individuals ofrare species and those with little foliage were sampled onlyonce for analysis in the laboratory where a quantitativeassay was used to test for cyanogenesis using freeze-dried ground tissue Because cyanogenesis is known tobe a polymorphic trait (eg Hughes 1991 Aikman et al1996) a minimum of three individuals of all species wastested more where species were common Owing to thediverse and heterogeneous nature of the forests multipleindividuals of each species were not always represented inthe plots Therefore for these rare species testing indivi-duals located outside the plots was required and still therewere several species for which only single samples wereobtained

A range of other factors was taken into considerationwhen sampling For example given that young leavesof tropical species in particular are typically more highlydefended than older leaves (Coley 1983 Coley and Barone1996) both young (soft expanding leaves) and old (recentfully expanded) leaves were tested for cyanogenesis wherepossible In addition depending on availability fruit andflowers from several species were tested Owing to the large

number of species and samples in the survey it was notpossible to examine seasonal trends in defence howeverindividuals of the majority of species were tested in wetand dry seasons for qualitative changes in cyanogenic sta-tus as there is some evidence for seasonal variation incyanogenesis (Seigler 1976b Janzen et al 1980 Kaplanet al 1983) Finally because edaphic factors have beenreported to affect the expression of cyanogenesis (egUrbanska 1982) species found on more than one substratetype were tested at each site

Sample collection handling and storage for quantitativeanalysis

In addition to samples for fresh cyanide tests whole leafsamples (again recent fully expanded leaves) were taken forquantification of cyanogenic glycosides All individuals ofspecies that produced a positive result and individuals ofrare species were sampled Depending on sampling condi-tions samples were either placed immediately into liquidnitrogen or placed in a sealed air-tight bag and kept on icefor 2ndash6 h until snap frozen in liquid nitrogen Frozensamples were transported to the laboratory on dry icefreeze-dried and stored on desiccant at 20 C for analysisFreeze-dried samples were ground using either a cooledIKA Labortechnic A10 Analytical Mill (Janke and KunkelStanfen Germany) or for smaller samples an Ultramat 2Dental Grinder (Southern Dental Industries Ltd BayswaterVictoria Australia)

Chemical analyses

Detection of cyanogenesis FeiglndashAnger papers Thepresence of cyanogenic compounds in fresh field sampleswas determined using FA indicator papers (Feigl andAnger 1966) FA papers were selected in preference topicrate papers because FA papers are more sensitive(Nahrstedt 1980) and less prone to giving false-positiveresults (Brinker and Seigler 1989) FA test papers wereprepared according to Brinker and Seigler (1989) BecauseFA papers can be sensitive to moisture and light (Brinkerand Seigler 1989) they were stored in the dark and ondesiccant until use

Fresh leaves (approx 1ndash2 g f wt) were crushed induplicate screw-top vials Old and young foliage sampleswere tested separately To facilitate cyanogenesis 05mL ofwater was added to one of the vials and pectinase fromRhizopus spp (Macerase Pectinase 441201 CalbiochemCalbiochem-Novabiochem Corp La Jolla CA USA)(04 g Lndash1) in 01 M TrisndashHCl (pH 68) was added to theother Pectinase has been found to have non-specific b-glyc-osidase activity (Brimer et al 1995) and therefore inthe absence of sufficient endogenous b-glycosidase enablestests for the presence of cyanogenic glycosides to be madeThe indicator papers were suspended above the tissue bymeans of the screw-top lid and vials were left at roomtemperature and checked after 12 and 24 h This 24 htime period used in other surveys (eg Dickenmann1982 Thomsen and Brimer 1997 Buhrmester et al2000 Lewis and Zona 2000) was selected to avoid spuri-ous test results due to substantial bacterial contamination

1020 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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which may occur beyond 24 h (Saupe et al 1982) Tissue ofknown cyanogenic species Prunus turneriana or Ryparosajavanica was used as a positive control Moderate tostrongly cyanogenic samples gave a positive result withina few minutes or up to a few hours while more weaklycyanogenic samples took several more hours In accordancewith the recommendations of Brinker and Seigler (1989) anew test was conducted for any samples producing a slowpositive response (24 h) in case of interference by microbialcyanogenesis In addition any samples with inconclusivecolour change were re-tested An individual was consideredcyanogenic if a positive repeatable result was obtainedand a species was considered cyanogenic if at leastone individual produced a consistent and repeatablepositive result A negative test result indicates the absenceof a cyanogenic glycoside or of the specific cyanogenicb-glycosidase or both

In the few instances where insufficient tissue was avail-able for both FA paper tests using fresh leaves and sub-sequent laboratory analysis cyanogenesis was determinedbased on the quantitative assay of freeze-dried groundleaf tissue (as described below) by comparison with anegative tissue control (eg Alstonia scholaris or Aglaiameridionalis)

In this study tests for cyanogenesis used approx1ndash2 g f wt of leaves which is larger than tissue samplestested in previous surveys (eg 50mg fwt by Lewis andZona 2000 and 200mg f wt by Thomsen and Brimer1997 Buhrmester et al 2000) According to Dickenmann(1982) who used 500mg f wt tissue a weak positive reac-tion with FA papers where part of the paper turns blueindicated approx 2ndash20mgHCNkgndash1 f wt while a strongreaction indicated gt50mgHCNkgndash1 f wt These lowervalues equate to just over 6ndash60mgHCNgndash1 d wt using aconversion based on the mean foliar water content ofseveral species in this study which was 70 In thisstudy based on fresh leaf tests and quantitative analysisof freeze-dried tissue from the same sample the thresholdsensitivity of FA papers was similar within the range5ndash8mgHCNgndash1 d wt This threshold sensitivity of FApapers corresponds well to the criteria of Adsersenet al (1988) for classifying individuals as cyanogenicwhere individuals with lt93 nmol HCNgndash1 f wt (approxim-ately equivalent to 8mgHCNgndash1 d wt) were consideredacyanogenic

Polymorphism and confirmation of acyanogenesis

For the purposes of the survey a species was consideredcyanogenic if at least one individual produced a repeatableand positive test result For a few species not all individualsproduced positive results using FA papers therefore somefurther investigation of the polymorphism for cyanogenesisin these species was conducted First to confirm the acy-anogenic character of the individuals producing negativeresults with FA papers a greater mass of freeze-dried tissue(100mg) was assayed quantitatively for the evolution ofcyanide (see below) and compared with a non-cyanogenicspecies A scholaris as a control for baseline noise in theassay Based on the criteria of Adsersen et al (1988) if

lt8mg HCNgndash1 d wt was detected then that individual wasconsidered acyanogenic

Quantification of cyanogenic glycosides

The concentration of cyanogenic glycoside in plant tis-sues was measured by trapping the cyanide (CN) liberatedfollowing hydrolysis of the glycoside in a well containing1 MNaOH (Brinker and Seigler 1989 Gleadow et al1998) Freeze-dried ground plant tissue (30ndash50mg) wasincubated for 20ndash24 h at 37 C with 1mL of 01M citratendashHCl (pH 55) Cyanide in the NaOH well was determinedusing the method of Gleadow and Woodrow (2002) adaptedfrom Brinker and Seigler (1989) for use with a photometricmicroplate reader (Labsystems Multiskan Ascent withincubator Labsystems Helsinki Finland) The method ishighly sensitive the determination of CN in NaOH isrelatively specific for cyanide and can detect as little as5mg Lndash1 (Brinker and Seigler 1989) The absorbance wasmeasured at 590 nm with NaCN as the standard

RESULTS

Survey for cyanogenesis

In total fresh leaf material from 401 woody plant speciesfrom 87 families was tested for cyanogenesis (Appendix)Cyanogenesis was found in 18 species from 13 familiesrepresenting 45 of species occurring within 1200m2 ofrainforest at each of five sites In each case the addition ofpectinase during incubation did not result in any additionalpositive results The test with added enzyme thereforeeffectively served as a replicate test for each sample Afurther two species were found to be cyanogenic butwere not included in the analysismdashthe non-woody Alocasiabrisbanensis and Helicia nortoniana which occurred out-side the plots (Table 1) The number of cyanogenic speciesat each site ranged from four to 10 The cyanogenic speciesranged in life form from climbing monocotyledons suchas the supplejack Flagellaria indica to 30m canopytrees such as the northern silky oak Cardwellia sublimisThe proportion of cyanogenic species ranged from 47 inupland rainforest at Mt Nomico to 65 in lowland rain-forest near Cape Tribulation Overall cyanogenic speciesaccounted for 73 of total basal area (range 12ndash134)The highest proportion was in highland rainforest on basaltsoil at Longlands Gap while forest on rhyolite in adjacentforest had the lowest proportion In lowland rainforestthe contribution of cyanogenic species to biomass wasalso high at 116 compared with 33 and 71 at MtNomico and Lamins Hill sites respectively

The cyanogenic species and concentrations of cyano-genic glycosides in leaves and other plant parts aresummarized in Table 1 The foliar concentrations of cyano-genic glycosides among species ranged from around8mgCNgndash1 d wtmdashthe concentration below which individu-als were considered acyanogenic based on the criteria ofAdsersen et al (1988) and the limits of FA paperdetectionmdashto very high concentrations in excess of severalmgCNgndash1 d wt (Table 1) For example fully expanded

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1021

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TABLE1

Su

mm

ary

of

the

con

cen

tra

tio

no

fcy

an

og

enic

gly

cosi

des

inle

ave

sa

nd

oth

erp

lan

tti

ssu

esfr

om

cya

no

gen

icsp

ecie

sfo

un

din

six

plo

tsin

up

lan

dh

igh

land

(U)

an

dlo

wla

nd

(L)

tro

pic

alr

ain

fore

sta

tfive

site

sh

igh

nu

trie

ntb

asa

ltsi

tes

atL

am

ins

Hil

l(B

1)

an

dL

on

gla

nd

sG

ap

(B2

)a

nd

low

nu

trie

nts

ites

on

gra

nit

ea

tMtN

om

ico

(G)

on

rhyo

lite

at

Lo

ng

lan

ds

Ga

p(R

)a

nd

on

met

am

orp

hic

sub

stra

ten

ear

Ca

pe

Tri

bu

lati

on

(M)

Concentrationofcyanogenic

glycosides

indifferentplantparts(m

gCNg1dw

t)

Form

Species

Fam

ily

Forest

Site

Leaf

Stem

Floralparts

Fruitseed

HA

loca

sia

bri

sba

nen

sis

(FM

Bailey)Domin

Araceae

UB1

ndashndash

ndashndash

TB

eils

chm

ied

iaco

llin

aBHyland

Lauraceae

UB1B2GR

Old28 4ndash1263(n

=36)

ndashndash

ndashYg1039ndash1391

TB

rom

bya

pla

tyn

emaFM

uell

Rutaceae

LM

156 4ndash1285(n

=20)

ndashndash

ndash0ndash10 5

(n=27)

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

UL

B1B1GRM

Old11 5ndash69 8

(n=21)

ndashBud779ndash851

Seedapprox8

Yg733 1tips219 1

Flwr130ndash334

TC

leis

tanth

us

myr

ian

thu

s(H

assk)Kurz

Euphorbiaceae

LM

Old1 1ndash17 3

(n=41)

ndashFlwr30 7

Mature160ndash377

Yg78 4ndash80 4

(n=3)

Immature821

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

UB1B2G

Old1325ndash4800(n

=6)

ndashBud733

ndashFlwr440ndash942

TE

laeo

carp

us

seri

cop

etalu

sFM

uell

Elaeocarpaceae

UGR

1100ndash5056(n

=10)

Sdlg1739ndash2679

ndash1028

Tree

135

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

UB1B2

10ndash81(n

=3)

ndashndash

ndashV

Fla

gel

lari

ain

dic

aL

Flagellariaceae

UL

B1GM

11ndash177 3

(n=6)

ndashndash

ndashT

Hel

icia

aust

rala

sica

FM

uell

Proteaceae

LM

42 2ndash155 2

(n=8)dagger

ndashndash

ndashT

Hel

icia

bla

keiForeman

Proteaceae

UB1

Mean17 9

(n=2)

ndashndash

ndashT

Hel

icia

no

rto

nia

na

(FM

Bailey)FM

Bailey

Proteaceae

LM

ndashndash

ndashndash

VP

ass

iflo

rasp(K

urandaBH12896)

Passifloraceae

LM

313ndash922(n

=4)

ndashndash

ndashT

Mis

cho

carp

us

exa

ng

ula

tus

(FM

uell)Radlk

Sapindaceae

UB1

Old133yg222(n

=1)

ndashndash

ndash

TM

isch

oca

rpu

sg

ran

dis

sim

us

(FM

uell)Radlk

Sapindaceae

UL

GM

49ndash680(n

=2)

761ndash2006(n

=4)z

ndashndash

ndash

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

UB1B2

Old10ndash208(n

=7)

ndashndash

ndashYg2000

VP

ars

on

sia

lati

foli

a(Benth)

STBlake

Apocynaceae

UB1GR

765ndash4835(n

=3)

ndashndash

ndash

TP

oly

scia

sa

ust

rali

an

a(FM

uell)Philipson

Araliaceae

UL

B1B2GRM

Oldlt6

8(n

=30)10ndash28 5

(n=16)

ndashndash

ndashYg10 8ndash29 6tips259

TP

runu

stu

rner

ian

a(FM

Bailey)Kalkman

Rosaceae

UL

B1M

old2472ndash4888(n

=4)

560ndash1100(n

=2)

ndashMature

seed400

Yg3128ndash6464Tip6000ndash8498

Rootapprox2000

Ygfruitseed8950

TR

ypa

rosa

java

nic

a(Blume)

Kurz

exKoordamp

Valeton

Achariaceae

LM

1749(n

=2)

ndashndash

Mature

seed5430

Yglf2560(n

=5)

Ygseed7760(n

=5)

Meansandranges

ofcyanogenicglycosides

concentrationaregiven

forfullyexpanded

(old)leaveswherenotspecifiedandforsomeyoung(yg)leavesfor

nindividualsLifeform

herb(H

)tree

(T)vine(V

)Cyanogenic

glycosideconcentrationsweremeasuredas

evolved

cyanidefollowinghydrolysisoffreeze-dried

tissuewithouttheadditionofnon-specificb-glycosidase(egem

ulsinpectinase)

A

loca

sia

bri

sba

nen

sis(a

herb)and

Hel

icia

nort

onia

nawhichoccurred

outsidetheplotswerenotincluded

intheanalysisnorsampledforquantitativeanalysis

Concentrationsofcyanogenic

glycosides

inseedflower

andyoungleaves

of

Ryp

aro

saja

van

icadetermined

byBLWebber

(unpubldata2004Webber

andWoodrow2004)

daggerOnly

oneindividual

of

Hel

icia

aust

rala

sica

occurred

within

theplotsseven

additional

saplingstreeswereopportunisticallytested

externalto

theplotsallwerecyanogenic

zOnly

oneindividual

of

Mis

cho

carp

us

gra

nd

issi

mu

soccurred

insideplotsat

MtNomicoadditional

individualsweresampledoutsideplotsincludingfourin

lowlandrainforestallwerecyanogenic

1022 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

leaves of numerous individuals of Polysciasaustraliana tested negative for cyanide and containedlt8mgCNgndash1 d wt while others tested positive and werein the range of 10ndash28 mgCNgndash1 d wt Even in the weaklycyanogenic samples only in very few cases did the colora-tion of the test paper change from 12 to 24 h in a qualit-ative sense Individuals of numerous species had highconcentrations of cyanogenic glycosides in excess of1mgCNgndash1 d wt (eg Mischocarpus grandissimusBrombya platynema and Beilschmiedia collina) and severalspecies contained extremely high concentrations of cyano-genic glycosides in mature tree leaves Notably fullyexpanded leaves from the tree species Elaeocarpus serico-petalus Clerodendrum grayi and Prunus turneriana hadconcentrations of cyanogenic glycosides up to 52 49and 48mgCNgndash1 d wt respectively (Table 1)

Qualitative and quantitative variation in cyanogenesis

Intra-population variation in cyanogenic glycosides Forthe majority of species initially identified as being cyano-genic all individuals tested were cyanogenic however theconcentrations of cyanogenic glycosides varied markedlybetween conspecific individuals (Table 1) For example allsaplings and trees of B collina at each site tested positivefor cyanogenesis but the concentration of cyanogenic glyc-osides in fully expanded leaves from saplings (n = 19)ranged from 232 to 12634mgCNgndash1 d wt at Mt Nomicoalone where B collina was most abundant In contrast tothis quantitative phenotypic variation in cyanogenesis con-specific individuals of a small number of species producedboth positive and negative FA paper test results Thisapparent qualitative polymorphism was most notable inB platynema a subcanopy tree species restricted to thelowland rainforest Within the population of B platynemaapprox 50 of individuals tested produced negativeFA test paper results for both young and old leaves andhad cyanogenic glycoside concentrations in the range06ndash68mgCNgndash1 d wt (Table 1) The addition ofbuffered pectinase during testing did not alter the qualitativeFA paper test result for any individual Among cyanogenicB platynema the concentrations of cyanogenic glyco-sides varied over orders of magnitude from 105 to12859mgCN gndash1 d wt (Table 1)

Negative results with FA papers were obtained for twoother species Cleistanthus myrianthus and Polysciasaustraliana Qualitative and quantitative analysis of oldleaves from individuals of these species indicated a highfrequency of acyanogenic individuals however unlikeB platynema the phenotypic expression of cyanogenesisvaried qualitatively with leaf age such that young leavesand leaf tips from these same individuals were consistentlycyanogenic aswere reproductive tissues fromC myrianthus(Table 1)

Even among less common species with smallsample sizes the concentrations of cyanogenic glycosidesvaried markedly between individuals For example inMischocarpus grandissimus the concentration amongsix individuals from two sites ranged from 49 to680mgCNgndash1 d wt at Mt Nomico and from 637 to

2006mgCNg1 d wt in lowland rainforest and amongthree individuals of the woody vine Parsonsia latifoliaconcentrations ranged from 765 to 4835 mgCNg1 d wtThere was no detectable affect of soil type on qualitativedetermination of cyanogenesis for individuals of the samespecies where cyanogenic species were found on differentsubstrates all individuals tested were cyanogenic

Intra-plant variation in cyanogenic glycoside concen-tration In addition to cyanogenic polymorphism both qual-itative and quantitative within populations of cyanogenicspecies the concentration of cyanogenic glycosides variedwith leaf age and plant part In all cases where tested youngleaves or leaf tips had higher concentrations of cyanogenicglycosides than older leaves (Table 1) In some cases theconcentrations in young and old leaves differed by over anorder of magnitude (eg C sublimis and Opisthiolepisheterophylla Table 1) In terms of determining the cyano-genic phenotype of an individual this difference betweenleaves of different ages was most notable for C myrianthusand P australiana Based on FA paper tests of mature fullyexpanded leaves of these species there were frequent acy-anogenic individuals however young leaves and leaf tipssampled from the same individuals of both species routinelytested positive for cyanogenesis Furthermore amongmature trees of C myrianthus which had low levels ofcyanogenic glycosides in old leaves (6ndash8mgCNg1 d wtie considered acyanogenic) the fruit in particularhad higher concentrations of cyanogenic glycosides(gt800mgCNg1 d wt Table 1) Overall few tests weremade of seeds or other reproductive tissues as part of thesurvey however where reproductive tissues were testedfrom species with cyanogenic leaves they were typicallycyanogenic and contained higher concentrations of cyano-genic glycosides although the concentration varied withthe maturity of the fruitseed For example the immatureseed and fruit (combined) from Prunus turneriana hadgt8mgCNg1 d wt compared with mature seed alone(lt1mgCNg1 d wt) while in R javanica cyanogenic gly-coside concentrations in flesh and seed of immature fruitdecreased with maturity (Table 1 Webber and Woodrow2004) One exception was the papery wind-dispersed seedsof C sublimis which yielded negligible cyanide (Table 1)

DISCUSSION

Cyanogenic species

Despite a substantial body of early work screening Austra-lian flora including some tropical flora for cyanogenesis(eg Petrie 1912 Smith and White 1918 Finnemore andCox 1928 Hurst 1942 Webb 1948 1949) few of thespecies tested in this study had previously been testedOf the 18 species from 13 families found to be cyanogenicin this study only one F indica has previously been repor-ted as cyanogenic (Petrie 1912) Thirteen of the cyanogenicspecies are endemic to Queensland or Australia several arerestricted to small areas within north east Queensland Forexample Clerodendrum grayi is found only in a small areaon the Atherton Tableland (Miller et al 2006a) while

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1023

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R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

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Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

which may occur beyond 24 h (Saupe et al 1982) Tissue ofknown cyanogenic species Prunus turneriana or Ryparosajavanica was used as a positive control Moderate tostrongly cyanogenic samples gave a positive result withina few minutes or up to a few hours while more weaklycyanogenic samples took several more hours In accordancewith the recommendations of Brinker and Seigler (1989) anew test was conducted for any samples producing a slowpositive response (24 h) in case of interference by microbialcyanogenesis In addition any samples with inconclusivecolour change were re-tested An individual was consideredcyanogenic if a positive repeatable result was obtainedand a species was considered cyanogenic if at leastone individual produced a consistent and repeatablepositive result A negative test result indicates the absenceof a cyanogenic glycoside or of the specific cyanogenicb-glycosidase or both

In the few instances where insufficient tissue was avail-able for both FA paper tests using fresh leaves and sub-sequent laboratory analysis cyanogenesis was determinedbased on the quantitative assay of freeze-dried groundleaf tissue (as described below) by comparison with anegative tissue control (eg Alstonia scholaris or Aglaiameridionalis)

In this study tests for cyanogenesis used approx1ndash2 g f wt of leaves which is larger than tissue samplestested in previous surveys (eg 50mg fwt by Lewis andZona 2000 and 200mg f wt by Thomsen and Brimer1997 Buhrmester et al 2000) According to Dickenmann(1982) who used 500mg f wt tissue a weak positive reac-tion with FA papers where part of the paper turns blueindicated approx 2ndash20mgHCNkgndash1 f wt while a strongreaction indicated gt50mgHCNkgndash1 f wt These lowervalues equate to just over 6ndash60mgHCNgndash1 d wt using aconversion based on the mean foliar water content ofseveral species in this study which was 70 In thisstudy based on fresh leaf tests and quantitative analysisof freeze-dried tissue from the same sample the thresholdsensitivity of FA papers was similar within the range5ndash8mgHCNgndash1 d wt This threshold sensitivity of FApapers corresponds well to the criteria of Adsersenet al (1988) for classifying individuals as cyanogenicwhere individuals with lt93 nmol HCNgndash1 f wt (approxim-ately equivalent to 8mgHCNgndash1 d wt) were consideredacyanogenic

Polymorphism and confirmation of acyanogenesis

For the purposes of the survey a species was consideredcyanogenic if at least one individual produced a repeatableand positive test result For a few species not all individualsproduced positive results using FA papers therefore somefurther investigation of the polymorphism for cyanogenesisin these species was conducted First to confirm the acy-anogenic character of the individuals producing negativeresults with FA papers a greater mass of freeze-dried tissue(100mg) was assayed quantitatively for the evolution ofcyanide (see below) and compared with a non-cyanogenicspecies A scholaris as a control for baseline noise in theassay Based on the criteria of Adsersen et al (1988) if

lt8mg HCNgndash1 d wt was detected then that individual wasconsidered acyanogenic

Quantification of cyanogenic glycosides

The concentration of cyanogenic glycoside in plant tis-sues was measured by trapping the cyanide (CN) liberatedfollowing hydrolysis of the glycoside in a well containing1 MNaOH (Brinker and Seigler 1989 Gleadow et al1998) Freeze-dried ground plant tissue (30ndash50mg) wasincubated for 20ndash24 h at 37 C with 1mL of 01M citratendashHCl (pH 55) Cyanide in the NaOH well was determinedusing the method of Gleadow and Woodrow (2002) adaptedfrom Brinker and Seigler (1989) for use with a photometricmicroplate reader (Labsystems Multiskan Ascent withincubator Labsystems Helsinki Finland) The method ishighly sensitive the determination of CN in NaOH isrelatively specific for cyanide and can detect as little as5mg Lndash1 (Brinker and Seigler 1989) The absorbance wasmeasured at 590 nm with NaCN as the standard

RESULTS

Survey for cyanogenesis

In total fresh leaf material from 401 woody plant speciesfrom 87 families was tested for cyanogenesis (Appendix)Cyanogenesis was found in 18 species from 13 familiesrepresenting 45 of species occurring within 1200m2 ofrainforest at each of five sites In each case the addition ofpectinase during incubation did not result in any additionalpositive results The test with added enzyme thereforeeffectively served as a replicate test for each sample Afurther two species were found to be cyanogenic butwere not included in the analysismdashthe non-woody Alocasiabrisbanensis and Helicia nortoniana which occurred out-side the plots (Table 1) The number of cyanogenic speciesat each site ranged from four to 10 The cyanogenic speciesranged in life form from climbing monocotyledons suchas the supplejack Flagellaria indica to 30m canopytrees such as the northern silky oak Cardwellia sublimisThe proportion of cyanogenic species ranged from 47 inupland rainforest at Mt Nomico to 65 in lowland rain-forest near Cape Tribulation Overall cyanogenic speciesaccounted for 73 of total basal area (range 12ndash134)The highest proportion was in highland rainforest on basaltsoil at Longlands Gap while forest on rhyolite in adjacentforest had the lowest proportion In lowland rainforestthe contribution of cyanogenic species to biomass wasalso high at 116 compared with 33 and 71 at MtNomico and Lamins Hill sites respectively

The cyanogenic species and concentrations of cyano-genic glycosides in leaves and other plant parts aresummarized in Table 1 The foliar concentrations of cyano-genic glycosides among species ranged from around8mgCNgndash1 d wtmdashthe concentration below which individu-als were considered acyanogenic based on the criteria ofAdsersen et al (1988) and the limits of FA paperdetectionmdashto very high concentrations in excess of severalmgCNgndash1 d wt (Table 1) For example fully expanded

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1021

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TABLE1

Su

mm

ary

of

the

con

cen

tra

tio

no

fcy

an

og

enic

gly

cosi

des

inle

ave

sa

nd

oth

erp

lan

tti

ssu

esfr

om

cya

no

gen

icsp

ecie

sfo

un

din

six

plo

tsin

up

lan

dh

igh

land

(U)

an

dlo

wla

nd

(L)

tro

pic

alr

ain

fore

sta

tfive

site

sh

igh

nu

trie

ntb

asa

ltsi

tes

atL

am

ins

Hil

l(B

1)

an

dL

on

gla

nd

sG

ap

(B2

)a

nd

low

nu

trie

nts

ites

on

gra

nit

ea

tMtN

om

ico

(G)

on

rhyo

lite

at

Lo

ng

lan

ds

Ga

p(R

)a

nd

on

met

am

orp

hic

sub

stra

ten

ear

Ca

pe

Tri

bu

lati

on

(M)

Concentrationofcyanogenic

glycosides

indifferentplantparts(m

gCNg1dw

t)

Form

Species

Fam

ily

Forest

Site

Leaf

Stem

Floralparts

Fruitseed

HA

loca

sia

bri

sba

nen

sis

(FM

Bailey)Domin

Araceae

UB1

ndashndash

ndashndash

TB

eils

chm

ied

iaco

llin

aBHyland

Lauraceae

UB1B2GR

Old28 4ndash1263(n

=36)

ndashndash

ndashYg1039ndash1391

TB

rom

bya

pla

tyn

emaFM

uell

Rutaceae

LM

156 4ndash1285(n

=20)

ndashndash

ndash0ndash10 5

(n=27)

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

UL

B1B1GRM

Old11 5ndash69 8

(n=21)

ndashBud779ndash851

Seedapprox8

Yg733 1tips219 1

Flwr130ndash334

TC

leis

tanth

us

myr

ian

thu

s(H

assk)Kurz

Euphorbiaceae

LM

Old1 1ndash17 3

(n=41)

ndashFlwr30 7

Mature160ndash377

Yg78 4ndash80 4

(n=3)

Immature821

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

UB1B2G

Old1325ndash4800(n

=6)

ndashBud733

ndashFlwr440ndash942

TE

laeo

carp

us

seri

cop

etalu

sFM

uell

Elaeocarpaceae

UGR

1100ndash5056(n

=10)

Sdlg1739ndash2679

ndash1028

Tree

135

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

UB1B2

10ndash81(n

=3)

ndashndash

ndashV

Fla

gel

lari

ain

dic

aL

Flagellariaceae

UL

B1GM

11ndash177 3

(n=6)

ndashndash

ndashT

Hel

icia

aust

rala

sica

FM

uell

Proteaceae

LM

42 2ndash155 2

(n=8)dagger

ndashndash

ndashT

Hel

icia

bla

keiForeman

Proteaceae

UB1

Mean17 9

(n=2)

ndashndash

ndashT

Hel

icia

no

rto

nia

na

(FM

Bailey)FM

Bailey

Proteaceae

LM

ndashndash

ndashndash

VP

ass

iflo

rasp(K

urandaBH12896)

Passifloraceae

LM

313ndash922(n

=4)

ndashndash

ndashT

Mis

cho

carp

us

exa

ng

ula

tus

(FM

uell)Radlk

Sapindaceae

UB1

Old133yg222(n

=1)

ndashndash

ndash

TM

isch

oca

rpu

sg

ran

dis

sim

us

(FM

uell)Radlk

Sapindaceae

UL

GM

49ndash680(n

=2)

761ndash2006(n

=4)z

ndashndash

ndash

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

UB1B2

Old10ndash208(n

=7)

ndashndash

ndashYg2000

VP

ars

on

sia

lati

foli

a(Benth)

STBlake

Apocynaceae

UB1GR

765ndash4835(n

=3)

ndashndash

ndash

TP

oly

scia

sa

ust

rali

an

a(FM

uell)Philipson

Araliaceae

UL

B1B2GRM

Oldlt6

8(n

=30)10ndash28 5

(n=16)

ndashndash

ndashYg10 8ndash29 6tips259

TP

runu

stu

rner

ian

a(FM

Bailey)Kalkman

Rosaceae

UL

B1M

old2472ndash4888(n

=4)

560ndash1100(n

=2)

ndashMature

seed400

Yg3128ndash6464Tip6000ndash8498

Rootapprox2000

Ygfruitseed8950

TR

ypa

rosa

java

nic

a(Blume)

Kurz

exKoordamp

Valeton

Achariaceae

LM

1749(n

=2)

ndashndash

Mature

seed5430

Yglf2560(n

=5)

Ygseed7760(n

=5)

Meansandranges

ofcyanogenicglycosides

concentrationaregiven

forfullyexpanded

(old)leaveswherenotspecifiedandforsomeyoung(yg)leavesfor

nindividualsLifeform

herb(H

)tree

(T)vine(V

)Cyanogenic

glycosideconcentrationsweremeasuredas

evolved

cyanidefollowinghydrolysisoffreeze-dried

tissuewithouttheadditionofnon-specificb-glycosidase(egem

ulsinpectinase)

A

loca

sia

bri

sba

nen

sis(a

herb)and

Hel

icia

nort

onia

nawhichoccurred

outsidetheplotswerenotincluded

intheanalysisnorsampledforquantitativeanalysis

Concentrationsofcyanogenic

glycosides

inseedflower

andyoungleaves

of

Ryp

aro

saja

van

icadetermined

byBLWebber

(unpubldata2004Webber

andWoodrow2004)

daggerOnly

oneindividual

of

Hel

icia

aust

rala

sica

occurred

within

theplotsseven

additional

saplingstreeswereopportunisticallytested

externalto

theplotsallwerecyanogenic

zOnly

oneindividual

of

Mis

cho

carp

us

gra

nd

issi

mu

soccurred

insideplotsat

MtNomicoadditional

individualsweresampledoutsideplotsincludingfourin

lowlandrainforestallwerecyanogenic

1022 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

leaves of numerous individuals of Polysciasaustraliana tested negative for cyanide and containedlt8mgCNgndash1 d wt while others tested positive and werein the range of 10ndash28 mgCNgndash1 d wt Even in the weaklycyanogenic samples only in very few cases did the colora-tion of the test paper change from 12 to 24 h in a qualit-ative sense Individuals of numerous species had highconcentrations of cyanogenic glycosides in excess of1mgCNgndash1 d wt (eg Mischocarpus grandissimusBrombya platynema and Beilschmiedia collina) and severalspecies contained extremely high concentrations of cyano-genic glycosides in mature tree leaves Notably fullyexpanded leaves from the tree species Elaeocarpus serico-petalus Clerodendrum grayi and Prunus turneriana hadconcentrations of cyanogenic glycosides up to 52 49and 48mgCNgndash1 d wt respectively (Table 1)

Qualitative and quantitative variation in cyanogenesis

Intra-population variation in cyanogenic glycosides Forthe majority of species initially identified as being cyano-genic all individuals tested were cyanogenic however theconcentrations of cyanogenic glycosides varied markedlybetween conspecific individuals (Table 1) For example allsaplings and trees of B collina at each site tested positivefor cyanogenesis but the concentration of cyanogenic glyc-osides in fully expanded leaves from saplings (n = 19)ranged from 232 to 12634mgCNgndash1 d wt at Mt Nomicoalone where B collina was most abundant In contrast tothis quantitative phenotypic variation in cyanogenesis con-specific individuals of a small number of species producedboth positive and negative FA paper test results Thisapparent qualitative polymorphism was most notable inB platynema a subcanopy tree species restricted to thelowland rainforest Within the population of B platynemaapprox 50 of individuals tested produced negativeFA test paper results for both young and old leaves andhad cyanogenic glycoside concentrations in the range06ndash68mgCNgndash1 d wt (Table 1) The addition ofbuffered pectinase during testing did not alter the qualitativeFA paper test result for any individual Among cyanogenicB platynema the concentrations of cyanogenic glyco-sides varied over orders of magnitude from 105 to12859mgCN gndash1 d wt (Table 1)

Negative results with FA papers were obtained for twoother species Cleistanthus myrianthus and Polysciasaustraliana Qualitative and quantitative analysis of oldleaves from individuals of these species indicated a highfrequency of acyanogenic individuals however unlikeB platynema the phenotypic expression of cyanogenesisvaried qualitatively with leaf age such that young leavesand leaf tips from these same individuals were consistentlycyanogenic aswere reproductive tissues fromC myrianthus(Table 1)

Even among less common species with smallsample sizes the concentrations of cyanogenic glycosidesvaried markedly between individuals For example inMischocarpus grandissimus the concentration amongsix individuals from two sites ranged from 49 to680mgCNgndash1 d wt at Mt Nomico and from 637 to

2006mgCNg1 d wt in lowland rainforest and amongthree individuals of the woody vine Parsonsia latifoliaconcentrations ranged from 765 to 4835 mgCNg1 d wtThere was no detectable affect of soil type on qualitativedetermination of cyanogenesis for individuals of the samespecies where cyanogenic species were found on differentsubstrates all individuals tested were cyanogenic

Intra-plant variation in cyanogenic glycoside concen-tration In addition to cyanogenic polymorphism both qual-itative and quantitative within populations of cyanogenicspecies the concentration of cyanogenic glycosides variedwith leaf age and plant part In all cases where tested youngleaves or leaf tips had higher concentrations of cyanogenicglycosides than older leaves (Table 1) In some cases theconcentrations in young and old leaves differed by over anorder of magnitude (eg C sublimis and Opisthiolepisheterophylla Table 1) In terms of determining the cyano-genic phenotype of an individual this difference betweenleaves of different ages was most notable for C myrianthusand P australiana Based on FA paper tests of mature fullyexpanded leaves of these species there were frequent acy-anogenic individuals however young leaves and leaf tipssampled from the same individuals of both species routinelytested positive for cyanogenesis Furthermore amongmature trees of C myrianthus which had low levels ofcyanogenic glycosides in old leaves (6ndash8mgCNg1 d wtie considered acyanogenic) the fruit in particularhad higher concentrations of cyanogenic glycosides(gt800mgCNg1 d wt Table 1) Overall few tests weremade of seeds or other reproductive tissues as part of thesurvey however where reproductive tissues were testedfrom species with cyanogenic leaves they were typicallycyanogenic and contained higher concentrations of cyano-genic glycosides although the concentration varied withthe maturity of the fruitseed For example the immatureseed and fruit (combined) from Prunus turneriana hadgt8mgCNg1 d wt compared with mature seed alone(lt1mgCNg1 d wt) while in R javanica cyanogenic gly-coside concentrations in flesh and seed of immature fruitdecreased with maturity (Table 1 Webber and Woodrow2004) One exception was the papery wind-dispersed seedsof C sublimis which yielded negligible cyanide (Table 1)

DISCUSSION

Cyanogenic species

Despite a substantial body of early work screening Austra-lian flora including some tropical flora for cyanogenesis(eg Petrie 1912 Smith and White 1918 Finnemore andCox 1928 Hurst 1942 Webb 1948 1949) few of thespecies tested in this study had previously been testedOf the 18 species from 13 families found to be cyanogenicin this study only one F indica has previously been repor-ted as cyanogenic (Petrie 1912) Thirteen of the cyanogenicspecies are endemic to Queensland or Australia several arerestricted to small areas within north east Queensland Forexample Clerodendrum grayi is found only in a small areaon the Atherton Tableland (Miller et al 2006a) while

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1023

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R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

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Dow

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Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

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TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TABLE1

Su

mm

ary

of

the

con

cen

tra

tio

no

fcy

an

og

enic

gly

cosi

des

inle

ave

sa

nd

oth

erp

lan

tti

ssu

esfr

om

cya

no

gen

icsp

ecie

sfo

un

din

six

plo

tsin

up

lan

dh

igh

land

(U)

an

dlo

wla

nd

(L)

tro

pic

alr

ain

fore

sta

tfive

site

sh

igh

nu

trie

ntb

asa

ltsi

tes

atL

am

ins

Hil

l(B

1)

an

dL

on

gla

nd

sG

ap

(B2

)a

nd

low

nu

trie

nts

ites

on

gra

nit

ea

tMtN

om

ico

(G)

on

rhyo

lite

at

Lo

ng

lan

ds

Ga

p(R

)a

nd

on

met

am

orp

hic

sub

stra

ten

ear

Ca

pe

Tri

bu

lati

on

(M)

Concentrationofcyanogenic

glycosides

indifferentplantparts(m

gCNg1dw

t)

Form

Species

Fam

ily

Forest

Site

Leaf

Stem

Floralparts

Fruitseed

HA

loca

sia

bri

sba

nen

sis

(FM

Bailey)Domin

Araceae

UB1

ndashndash

ndashndash

TB

eils

chm

ied

iaco

llin

aBHyland

Lauraceae

UB1B2GR

Old28 4ndash1263(n

=36)

ndashndash

ndashYg1039ndash1391

TB

rom

bya

pla

tyn

emaFM

uell

Rutaceae

LM

156 4ndash1285(n

=20)

ndashndash

ndash0ndash10 5

(n=27)

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

UL

B1B1GRM

Old11 5ndash69 8

(n=21)

ndashBud779ndash851

Seedapprox8

Yg733 1tips219 1

Flwr130ndash334

TC

leis

tanth

us

myr

ian

thu

s(H

assk)Kurz

Euphorbiaceae

LM

Old1 1ndash17 3

(n=41)

ndashFlwr30 7

Mature160ndash377

Yg78 4ndash80 4

(n=3)

Immature821

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

UB1B2G

Old1325ndash4800(n

=6)

ndashBud733

ndashFlwr440ndash942

TE

laeo

carp

us

seri

cop

etalu

sFM

uell

Elaeocarpaceae

UGR

1100ndash5056(n

=10)

Sdlg1739ndash2679

ndash1028

Tree

135

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

UB1B2

10ndash81(n

=3)

ndashndash

ndashV

Fla

gel

lari

ain

dic

aL

Flagellariaceae

UL

B1GM

11ndash177 3

(n=6)

ndashndash

ndashT

Hel

icia

aust

rala

sica

FM

uell

Proteaceae

LM

42 2ndash155 2

(n=8)dagger

ndashndash

ndashT

Hel

icia

bla

keiForeman

Proteaceae

UB1

Mean17 9

(n=2)

ndashndash

ndashT

Hel

icia

no

rto

nia

na

(FM

Bailey)FM

Bailey

Proteaceae

LM

ndashndash

ndashndash

VP

ass

iflo

rasp(K

urandaBH12896)

Passifloraceae

LM

313ndash922(n

=4)

ndashndash

ndashT

Mis

cho

carp

us

exa

ng

ula

tus

(FM

uell)Radlk

Sapindaceae

UB1

Old133yg222(n

=1)

ndashndash

ndash

TM

isch

oca

rpu

sg

ran

dis

sim

us

(FM

uell)Radlk

Sapindaceae

UL

GM

49ndash680(n

=2)

761ndash2006(n

=4)z

ndashndash

ndash

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

UB1B2

Old10ndash208(n

=7)

ndashndash

ndashYg2000

VP

ars

on

sia

lati

foli

a(Benth)

STBlake

Apocynaceae

UB1GR

765ndash4835(n

=3)

ndashndash

ndash

TP

oly

scia

sa

ust

rali

an

a(FM

uell)Philipson

Araliaceae

UL

B1B2GRM

Oldlt6

8(n

=30)10ndash28 5

(n=16)

ndashndash

ndashYg10 8ndash29 6tips259

TP

runu

stu

rner

ian

a(FM

Bailey)Kalkman

Rosaceae

UL

B1M

old2472ndash4888(n

=4)

560ndash1100(n

=2)

ndashMature

seed400

Yg3128ndash6464Tip6000ndash8498

Rootapprox2000

Ygfruitseed8950

TR

ypa

rosa

java

nic

a(Blume)

Kurz

exKoordamp

Valeton

Achariaceae

LM

1749(n

=2)

ndashndash

Mature

seed5430

Yglf2560(n

=5)

Ygseed7760(n

=5)

Meansandranges

ofcyanogenicglycosides

concentrationaregiven

forfullyexpanded

(old)leaveswherenotspecifiedandforsomeyoung(yg)leavesfor

nindividualsLifeform

herb(H

)tree

(T)vine(V

)Cyanogenic

glycosideconcentrationsweremeasuredas

evolved

cyanidefollowinghydrolysisoffreeze-dried

tissuewithouttheadditionofnon-specificb-glycosidase(egem

ulsinpectinase)

A

loca

sia

bri

sba

nen

sis(a

herb)and

Hel

icia

nort

onia

nawhichoccurred

outsidetheplotswerenotincluded

intheanalysisnorsampledforquantitativeanalysis

Concentrationsofcyanogenic

glycosides

inseedflower

andyoungleaves

of

Ryp

aro

saja

van

icadetermined

byBLWebber

(unpubldata2004Webber

andWoodrow2004)

daggerOnly

oneindividual

of

Hel

icia

aust

rala

sica

occurred

within

theplotsseven

additional

saplingstreeswereopportunisticallytested

externalto

theplotsallwerecyanogenic

zOnly

oneindividual

of

Mis

cho

carp

us

gra

nd

issi

mu

soccurred

insideplotsat

MtNomicoadditional

individualsweresampledoutsideplotsincludingfourin

lowlandrainforestallwerecyanogenic

1022 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

leaves of numerous individuals of Polysciasaustraliana tested negative for cyanide and containedlt8mgCNgndash1 d wt while others tested positive and werein the range of 10ndash28 mgCNgndash1 d wt Even in the weaklycyanogenic samples only in very few cases did the colora-tion of the test paper change from 12 to 24 h in a qualit-ative sense Individuals of numerous species had highconcentrations of cyanogenic glycosides in excess of1mgCNgndash1 d wt (eg Mischocarpus grandissimusBrombya platynema and Beilschmiedia collina) and severalspecies contained extremely high concentrations of cyano-genic glycosides in mature tree leaves Notably fullyexpanded leaves from the tree species Elaeocarpus serico-petalus Clerodendrum grayi and Prunus turneriana hadconcentrations of cyanogenic glycosides up to 52 49and 48mgCNgndash1 d wt respectively (Table 1)

Qualitative and quantitative variation in cyanogenesis

Intra-population variation in cyanogenic glycosides Forthe majority of species initially identified as being cyano-genic all individuals tested were cyanogenic however theconcentrations of cyanogenic glycosides varied markedlybetween conspecific individuals (Table 1) For example allsaplings and trees of B collina at each site tested positivefor cyanogenesis but the concentration of cyanogenic glyc-osides in fully expanded leaves from saplings (n = 19)ranged from 232 to 12634mgCNgndash1 d wt at Mt Nomicoalone where B collina was most abundant In contrast tothis quantitative phenotypic variation in cyanogenesis con-specific individuals of a small number of species producedboth positive and negative FA paper test results Thisapparent qualitative polymorphism was most notable inB platynema a subcanopy tree species restricted to thelowland rainforest Within the population of B platynemaapprox 50 of individuals tested produced negativeFA test paper results for both young and old leaves andhad cyanogenic glycoside concentrations in the range06ndash68mgCNgndash1 d wt (Table 1) The addition ofbuffered pectinase during testing did not alter the qualitativeFA paper test result for any individual Among cyanogenicB platynema the concentrations of cyanogenic glyco-sides varied over orders of magnitude from 105 to12859mgCN gndash1 d wt (Table 1)

Negative results with FA papers were obtained for twoother species Cleistanthus myrianthus and Polysciasaustraliana Qualitative and quantitative analysis of oldleaves from individuals of these species indicated a highfrequency of acyanogenic individuals however unlikeB platynema the phenotypic expression of cyanogenesisvaried qualitatively with leaf age such that young leavesand leaf tips from these same individuals were consistentlycyanogenic aswere reproductive tissues fromC myrianthus(Table 1)

Even among less common species with smallsample sizes the concentrations of cyanogenic glycosidesvaried markedly between individuals For example inMischocarpus grandissimus the concentration amongsix individuals from two sites ranged from 49 to680mgCNgndash1 d wt at Mt Nomico and from 637 to

2006mgCNg1 d wt in lowland rainforest and amongthree individuals of the woody vine Parsonsia latifoliaconcentrations ranged from 765 to 4835 mgCNg1 d wtThere was no detectable affect of soil type on qualitativedetermination of cyanogenesis for individuals of the samespecies where cyanogenic species were found on differentsubstrates all individuals tested were cyanogenic

Intra-plant variation in cyanogenic glycoside concen-tration In addition to cyanogenic polymorphism both qual-itative and quantitative within populations of cyanogenicspecies the concentration of cyanogenic glycosides variedwith leaf age and plant part In all cases where tested youngleaves or leaf tips had higher concentrations of cyanogenicglycosides than older leaves (Table 1) In some cases theconcentrations in young and old leaves differed by over anorder of magnitude (eg C sublimis and Opisthiolepisheterophylla Table 1) In terms of determining the cyano-genic phenotype of an individual this difference betweenleaves of different ages was most notable for C myrianthusand P australiana Based on FA paper tests of mature fullyexpanded leaves of these species there were frequent acy-anogenic individuals however young leaves and leaf tipssampled from the same individuals of both species routinelytested positive for cyanogenesis Furthermore amongmature trees of C myrianthus which had low levels ofcyanogenic glycosides in old leaves (6ndash8mgCNg1 d wtie considered acyanogenic) the fruit in particularhad higher concentrations of cyanogenic glycosides(gt800mgCNg1 d wt Table 1) Overall few tests weremade of seeds or other reproductive tissues as part of thesurvey however where reproductive tissues were testedfrom species with cyanogenic leaves they were typicallycyanogenic and contained higher concentrations of cyano-genic glycosides although the concentration varied withthe maturity of the fruitseed For example the immatureseed and fruit (combined) from Prunus turneriana hadgt8mgCNg1 d wt compared with mature seed alone(lt1mgCNg1 d wt) while in R javanica cyanogenic gly-coside concentrations in flesh and seed of immature fruitdecreased with maturity (Table 1 Webber and Woodrow2004) One exception was the papery wind-dispersed seedsof C sublimis which yielded negligible cyanide (Table 1)

DISCUSSION

Cyanogenic species

Despite a substantial body of early work screening Austra-lian flora including some tropical flora for cyanogenesis(eg Petrie 1912 Smith and White 1918 Finnemore andCox 1928 Hurst 1942 Webb 1948 1949) few of thespecies tested in this study had previously been testedOf the 18 species from 13 families found to be cyanogenicin this study only one F indica has previously been repor-ted as cyanogenic (Petrie 1912) Thirteen of the cyanogenicspecies are endemic to Queensland or Australia several arerestricted to small areas within north east Queensland Forexample Clerodendrum grayi is found only in a small areaon the Atherton Tableland (Miller et al 2006a) while

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1023

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R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

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53

13)Rubiaceae

(3)

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on

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uell)Gibbs

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917

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(2)

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da

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iosa

(FM

Bailey)Domin

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(2)

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1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

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uell)Radlk

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ValetonD

Achariaceae

dagger++

LM

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mel

(1911)

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caes

ia+C

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(1919)

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(2)

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(CTW

hite)

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ctaRadlk

Sapindaceae

(3)

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tCarbine

LW

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sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

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nto

ph

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TS

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hn

soniiFM

uell

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naceae

(5)

UB1

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whit

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Cunoniaceae

(1)

UG

TS

colo

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ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

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on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

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Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

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VS

mil

ax

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cip

hyl

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Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

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So

lan

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sp

Solanaceae

(2)

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viri

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Dunal

Solanaceae

(1)

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eno

stem

on

lob

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s(FM

uell)LSSm

Sphenostem

onaceae

(3)

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MELU102111

SHT

Ste

ga

nth

era

au

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lia

naCTW

hite

Monim

iaceae

(2)

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era

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coo

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(FM

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iaceae

(4)

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TSte

noca

rpus

reti

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hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

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Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

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nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

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rali

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s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

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min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

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Symplocaceae

(2)

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loco

sco

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Symplocaceae

(2)

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Noot

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(3)

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SH

Sym

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hiteamp

WDFrancis

Symplocaceae

(3)

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MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

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Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

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MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

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MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

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son

ii(FM

uell)BHyland

Myrtaceae

(3)

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nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

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ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

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yzyg

ium

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no

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mu

mCraven

Myrtaceae

(1)

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pyr

ace

um

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Myrtaceae

(3)

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m(CTW

hite)

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(1)

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yzyg

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wes

aBHyland

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(3)

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102249

TS

yzyg

ium

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son

iisubsp

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biu

m(FM

uell)BHyland

Myrtaceae

(3)

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SH

Taber

naem

onta

na

pandaca

quiLam

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(1)

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Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

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(4)

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Ta

sma

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iain

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Winteraceae

(3)dry

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ernst

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erry

i(FM

Bailey)

Merrex

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(2)

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no

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avar

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aFM

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(2)

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an

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ra(Benth)JRPerkins

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iaceae

(6)

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(DaviesCreek

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daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

leaves of numerous individuals of Polysciasaustraliana tested negative for cyanide and containedlt8mgCNgndash1 d wt while others tested positive and werein the range of 10ndash28 mgCNgndash1 d wt Even in the weaklycyanogenic samples only in very few cases did the colora-tion of the test paper change from 12 to 24 h in a qualit-ative sense Individuals of numerous species had highconcentrations of cyanogenic glycosides in excess of1mgCNgndash1 d wt (eg Mischocarpus grandissimusBrombya platynema and Beilschmiedia collina) and severalspecies contained extremely high concentrations of cyano-genic glycosides in mature tree leaves Notably fullyexpanded leaves from the tree species Elaeocarpus serico-petalus Clerodendrum grayi and Prunus turneriana hadconcentrations of cyanogenic glycosides up to 52 49and 48mgCNgndash1 d wt respectively (Table 1)

Qualitative and quantitative variation in cyanogenesis

Intra-population variation in cyanogenic glycosides Forthe majority of species initially identified as being cyano-genic all individuals tested were cyanogenic however theconcentrations of cyanogenic glycosides varied markedlybetween conspecific individuals (Table 1) For example allsaplings and trees of B collina at each site tested positivefor cyanogenesis but the concentration of cyanogenic glyc-osides in fully expanded leaves from saplings (n = 19)ranged from 232 to 12634mgCNgndash1 d wt at Mt Nomicoalone where B collina was most abundant In contrast tothis quantitative phenotypic variation in cyanogenesis con-specific individuals of a small number of species producedboth positive and negative FA paper test results Thisapparent qualitative polymorphism was most notable inB platynema a subcanopy tree species restricted to thelowland rainforest Within the population of B platynemaapprox 50 of individuals tested produced negativeFA test paper results for both young and old leaves andhad cyanogenic glycoside concentrations in the range06ndash68mgCNgndash1 d wt (Table 1) The addition ofbuffered pectinase during testing did not alter the qualitativeFA paper test result for any individual Among cyanogenicB platynema the concentrations of cyanogenic glyco-sides varied over orders of magnitude from 105 to12859mgCN gndash1 d wt (Table 1)

Negative results with FA papers were obtained for twoother species Cleistanthus myrianthus and Polysciasaustraliana Qualitative and quantitative analysis of oldleaves from individuals of these species indicated a highfrequency of acyanogenic individuals however unlikeB platynema the phenotypic expression of cyanogenesisvaried qualitatively with leaf age such that young leavesand leaf tips from these same individuals were consistentlycyanogenic aswere reproductive tissues fromC myrianthus(Table 1)

Even among less common species with smallsample sizes the concentrations of cyanogenic glycosidesvaried markedly between individuals For example inMischocarpus grandissimus the concentration amongsix individuals from two sites ranged from 49 to680mgCNgndash1 d wt at Mt Nomico and from 637 to

2006mgCNg1 d wt in lowland rainforest and amongthree individuals of the woody vine Parsonsia latifoliaconcentrations ranged from 765 to 4835 mgCNg1 d wtThere was no detectable affect of soil type on qualitativedetermination of cyanogenesis for individuals of the samespecies where cyanogenic species were found on differentsubstrates all individuals tested were cyanogenic

Intra-plant variation in cyanogenic glycoside concen-tration In addition to cyanogenic polymorphism both qual-itative and quantitative within populations of cyanogenicspecies the concentration of cyanogenic glycosides variedwith leaf age and plant part In all cases where tested youngleaves or leaf tips had higher concentrations of cyanogenicglycosides than older leaves (Table 1) In some cases theconcentrations in young and old leaves differed by over anorder of magnitude (eg C sublimis and Opisthiolepisheterophylla Table 1) In terms of determining the cyano-genic phenotype of an individual this difference betweenleaves of different ages was most notable for C myrianthusand P australiana Based on FA paper tests of mature fullyexpanded leaves of these species there were frequent acy-anogenic individuals however young leaves and leaf tipssampled from the same individuals of both species routinelytested positive for cyanogenesis Furthermore amongmature trees of C myrianthus which had low levels ofcyanogenic glycosides in old leaves (6ndash8mgCNg1 d wtie considered acyanogenic) the fruit in particularhad higher concentrations of cyanogenic glycosides(gt800mgCNg1 d wt Table 1) Overall few tests weremade of seeds or other reproductive tissues as part of thesurvey however where reproductive tissues were testedfrom species with cyanogenic leaves they were typicallycyanogenic and contained higher concentrations of cyano-genic glycosides although the concentration varied withthe maturity of the fruitseed For example the immatureseed and fruit (combined) from Prunus turneriana hadgt8mgCNg1 d wt compared with mature seed alone(lt1mgCNg1 d wt) while in R javanica cyanogenic gly-coside concentrations in flesh and seed of immature fruitdecreased with maturity (Table 1 Webber and Woodrow2004) One exception was the papery wind-dispersed seedsof C sublimis which yielded negligible cyanide (Table 1)

DISCUSSION

Cyanogenic species

Despite a substantial body of early work screening Austra-lian flora including some tropical flora for cyanogenesis(eg Petrie 1912 Smith and White 1918 Finnemore andCox 1928 Hurst 1942 Webb 1948 1949) few of thespecies tested in this study had previously been testedOf the 18 species from 13 families found to be cyanogenicin this study only one F indica has previously been repor-ted as cyanogenic (Petrie 1912) Thirteen of the cyanogenicspecies are endemic to Queensland or Australia several arerestricted to small areas within north east Queensland Forexample Clerodendrum grayi is found only in a small areaon the Atherton Tableland (Miller et al 2006a) while

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1023

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R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

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plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

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Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

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Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

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Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

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Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

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Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

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Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

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Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

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Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

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Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

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Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

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Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

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Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

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Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

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Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

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van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

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Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

R javanica occurs in a limited area within lowland rain-forest north of the Daintree River Cyanogenesis is reportedfor the first time in the genera Beilschmiedia CardwelliaCleistanthus Elaeocarpus Embelia MischocarpusOpisthiolepis Parsonsia and Polyscias The number ofnew reports at the generic level may in part reflect thehigh level of endemism among the Australian tropical rain-forest flora In addition several species are from families inwhich cyanogenesis has been rarely reported if at all Forexample cyanogenesis is rare in ElaeocarpaceaeLauraceae Apocynaceae Myrsinaceae and Araliaceaefamilies Following are descriptions of each cyanogenicspecies (listed alphabetically by family) discussed in rela-tion to previous reports of cyanogenesis within the relevanttaxonomic groups

Ryparosa javanica (Blume) Kurz ex Koord ampValeton (Achariaceae)

Ryparosa javanica is currently the subject of taxonomicrevision (Webber 2005) The Queensland Ryparosa spcurrently known as R javanica is endemic to lowland rain-forest north of the Daintree River to Cape Tribulation northeast Queensland This species was found to be cyanogenicearly in this survey and has subsequently been the focusof extensive population-level studies of cyanogenesis(Webber 2005) Reports of cyanogenesis in this genusare common for example R javanica (sensu stricto) andR caesia are cyanogenic (Rosenthaler 1919) Cyanogen-esis has been reported among other genera in Achariaceaeeg Hydnocarpus Calancoba Ceratiosicyos GynocardiaErythrospermum Pangium and Kiggelaria (Rosenthaler1919 Tjon Sie Fat 1979a Jensen and Nielsen 1986)Many of these genera were previously in the Flacourtiaceaeuntil recent revisions saw most cyanogenic genera assignedto the Achariaceae (Chase et al 2002) including the tribePangieae of which Ryparosa is a member Cyanogenesiswas considered a useful taxonomic marker in Flacourtiaceae(Spencer and Seigler 1985) this utility being apparent inthe revision of the family (Chase et al 2002) All individu-als in sizeable populations of R javanica were cyanogenicand the cyanogenic glycoside in R javanica has beenidentified as gynocardin (Webber 2005) a cyclopentenoidcyanogenic glycoside typical of Achariaceae (Jaroszewskiand Olafsdottir 1987) Cyanogenic glycosides derivedfrom valineisoleucine have also been reported inspecies formerly in the Flacourtiaceae (Lechtenberg andNahrstedt 1999)

Parsonsia latifolia (Benth) STBlake (Apocynaceae)

This is the first report of cyanogenesis in the genusParsonsia a genus of woody or semi-woody climbers(130 species) distributed from Southeast Asia to AustraliaNew Caledonia and New Zealand Members of the Apo-cynaceae family (220 genera 2000 species) commonly haveclear or milky latex and frequently contain alkaloids(Mabberley 1990 Collins et al 1990) Parsonia latifolia(diameter up to 9 cm) has milky white latex and is endemicto Australia (Forster and Williams 1996) It is found inlowland and highland rainforest in noth east Queensland

parts of New South Wales and the Northern Territory(Hyland et al 2003) Reports of cyanogenesis in Apocyn-aceae and the order Gentianales are few Gibbs (1974) whoreported negative results for Parsonsia eucalyptifolia andP lanceolata found only Alstonia scholaris (L) RBr to becyanogenic and noted positive reports for four otherspecies Tests for cyanogenesis in the Apocynaceae arehowever apparently limited with negative reports foronly approx 20 species (Gibbs 1974 Adsersen et al1988 Thomsen and Brimer 1997) In this study leafsamples of all ages from multiple A scholaris trees andseedlings gave negative FA paper results quantitativeassaying confirmed the absence of cyanogenesis No cyano-genic constituents in the family have been characterized

Polyscias australiana (FMuell) Philipson (Araliaceae)

This is the first report of cyanogenesis in the genusPolyscias a genus of approx 100 species distributedthroughout Africa Asia Malesia Australia and the PacificIslands In addition cyanogenesis is very rare in the orderUmbellales Gibbs (1974) reported only negative results orsome doubtful positive results in a few members ofAraliaceae (Nothopanax sp and Schefflera sp) and theUmbelliferae Subsequently only Aralia spinosa L(above-ground parts) has been reported as cyanogenic(Seigler 1976b) Polyscias australiana is often considereda regrowth species in disturbed rainforest and commonlyoccurs at rainforest margins Concentrations of cyanogenicglycosides in this species were variable and in matureleaves were commonly less than the threshold value usedfor classifying individuals as cyanogenic in this study(lt8mgCNgndash1 dwt) however the species was consideredcyanogenic on the basis of repeatable positive results withFA papers for multiple individuals in particular whenanalysing young foliage Interestingly in reporting cyan-ogenesis in A spinosa Seigler (1976b) noted substantialseasonal variation in that the individual was cyanogenic atonly one of three testing times throughout the year Analysisof the distribution of cyanogenic glycosides in fruitflowers and seasonal variation in foliar concentrations inP australiana would better characterize cyanogenesisin this species

Elaeocarpus sericopetalus FMuell (Elaeocarpaceae)

Elaeocarpus sericopetalus (lsquoNorthern Quandongrsquo) is asmall to medium sized tree endemic to tropical rainforestwithin the Cook and Kennedy Districts of north easternQueensland This is the first report of cyanogenesis inthe genus Elaeocarpus Elaeocarpus sericopetalus wasthe only cyanogenic species identified among eightElaeocarpus spp and 11 species from the Elaeocarpaceaefamily tested in this study (Appendix) The extremely highfoliar concentrations of cyanogenic glycosides (up to52mgCNgndash1 d wt in mature field-grown leaves) inE sericopetalus rank it among the most cyanogenic treespecies ever reported Cyanogenesis is rare within thefamily Elaeocarpaceae and the order Malvales (Gibbs1974 Hegnauer 1990 Lechtenberg and Nahrstedt1999) whereas alkaloids (eg indolizidine alkaloids) are

1024 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

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Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

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Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

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Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

considered common (Gibbs 1974 Hegnauer 1990Mabberley 1990) In the Elaeocarpaceae family cyanogen-esis has previously been reported in only two species theleaves of Vallea stipularis lsquopyrifoliarsquo FBallard (Gibbs1974) and the leaves [Greshoff (1898) cited in Hegnauer(1973)] and the bark (Pammel 1911 Rosenthaler 1919) ofSloanea sigun (Blume) KSchum (syn Echinocarpussigun) were found to be cyanogenic Sambunigrin wasisolated from the leaves of S sigun [R Hegnauer andL H Fikenscher unpubl data (1983) cited in Hegnauer(1990)] the only previous report of a cyanogenic constitu-ent in Elaeocarpaceae and Malvales Characterization of theprincipal foliar cyanogenic glycosidemdashan apparentlyunusual phenylalanine-derived glycoside with an organicacid residuemdashis ongoing (Miller 2004)

Cleistanthus myrianthus (Hassk) Kurz (Euphorbiaceae)

This appears to be the first report of cyanogenesis inthe genus Cleistanthus which consists of 100ndash140 speciesCleistanthus myrianthus is a subcanopy rainforest tree (to7m) found in the lowland and foothills from the DaintreeRiver to Rossville north east Queensland (Cooper andCooper 1994) In Australia it is classified as rare on thebasis of this limited distribution but it is also found inSoutheast Asia and Malesia (Hyland et al 2003) Cyano-genesis is especially common in Euphorbiaceae (300genera 7500 species) and is found in many genera includ-ing Bridelia Euphorbia and Phyllanthus as well as theeconomically important species Hevea brasiliensis (rubbertree) and Manihot esculenta (cassava) (eg Rosenthaler1919 Tjon Sie Fat 1979a Seigler et al 1979 Adsersenet al 1988)

The chemotaxonomic utility of cyanogenesis as wellas other secondary metabolites (eg alkaloids and terpenes)has been demonstrated in Euphorbiaceae (Seigler 1994)Cyanogenic glycosides are useful at the infra-familiallevel in the Euphorbiaceae (van Valen 1978 Seigler1994) the species in the subfamily Phyllanthoideae typic-ally contain the tyrosine-derived cyanogenic glycosidesdhurrin taxiphyllin or triglochinin while species in thesubfamily Crotonoideae (sensu Pax and Hoffman 1931see van Valen 1978) including Hevea and Manihot pro-duce cyanogenic glycosides derived from valine and iso-leucine (eg linamarin and lotaustralin) (van Valen 1978Nahrstedt 1987 Seigler 1994) Given this pattern one maypredict Cleistanthusmdashassigned to the Phyllanthoideaemdashtocontain cyanogens derived from tyrosine

Flagellaria indica L (Flagellariaceae)

Flagellaria indica (lsquothe supplejackrsquo) a leaf tendril clim-ber with cane-like stems is known to be cyanogenic (Petrie1912 Webb 1948 Gibbs 1974 Morley and Toelken1983) Young shoots were suspected of poisoning stockin Australia (Everist 1981) and it has also been reportedto have medicinal properties (eg tumour inhibition Collinset al 1990) In other countries it has been used as a hairwash and as a contraceptive (see Hyland et al 2003)but was not apparently used medicinally by aboriginalAustralians (Morley and Toelken 1983) Interestingly

monocotyledons are typically characterized by cyanogenicglycosides biosynthetically derived from tyrosine(Lechtenberg and Nahrstedt 1999) Consistent with thisthe tyrosine-derived cyanogenic glycoside triglochininwas isolated from the stem (and rhizome) of F indica(L H Fikenscher unpubl data cited in Hegnauer 1966)although meta-hydroxylated valine or isoleucine andleucine-derived glycosides have also been found inmonocotyledons (Nahrstedt 1987 Lechtenberg andNahrstedt 1999)

Clerodendrum grayi Munir (Lamiaceae)

Clerodendrum grayi is a rare subcanopy tree endemic tothe northern part of Queensland Australia (Munir 1989)Recent revisions of the division between Lamiaceae andVerbenaceae families (Cantino 1992 Wagstaff et al1997 1998) transferred the genus Clerodendrum and othershistorically in the Verbenaceae family to the Lamiaceaefamily Cyanogenesis in Lamiaceae and also Verbenaceaehas rarely been reported Even within the order Lamialescyanogenesis is considered rare (Gibbs 1974) In theLamiaceae known for its culinary and medicinal herbs[eg Lavandula (lavender) Mentha (mint)] typical con-stituents are monoterpenoids diterpenes or triterpenes aswell as flavonoids and iridoid glycosides (Gibbs 1974Hegnauer 1989 Taskova et al 1997) In a survey of theflora of the Galapagos Islands Clerodendrum molle varmolle was found to be cyanogenic (Adsersen et al 1988see also Gibbs 1974 Tjon sie Fat 1979a) In additionseveral species of Clerodendrum are known to be toxic(Hurst 1942 Webb 1948 CFSAN 2003) however thepoison is not detailed

In this study the extremely high foliar concentrations ofcyanogenic compoundsmdashup to 48mgCNg1 d wt inmature field-grown tree leavesmdashare among the highestreported for tree leaves (Table 1) Two cyanogenic glycos-ides were purified from the leaf tissue of C grayi (Milleret al 2006a) Prunasin and its primerveroside the rarediglycoside lucumin (Eyjolfsson 1971) were found inthe ratio 1 158 (molmol) (Miller et al 2006a) the firstreported co-occurrence of these glycosides and the firstconfirmed report of lucumin in vegetative tissue (seeThomsen and Brimer 1997) Given the relatively rarityof reports of cyanogenic glycosides from the Lamiaceaeand even within the order Lamiales it is difficult to drawany conclusions about the biogenetic origins of glycosideswithin these taxonomic groups Refer to Miller et al(2006a) for a detailed discussion of cyanogenesis inC grayi and associated taxa

Beilschmiedia collina BHyland (Lauraceae)

Beilschmiedia collina (lsquothe mountain blush walnutrsquo) is atree species endemic to Queensland rainforest (Cooper andCooper 1994) Cyanogenesis is very rare in Lauraceae(Gibbs 1974 Hegnauer 1989) having only been reportedfrom Cinnamomum camphora and Litsea glutinosa (Gibbs1974) with one other species (Nectranda megapotamica)reported to have cyanogenic glycosides but apparentlylacks the catabolic enzymes requiring further investigation

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1025

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(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

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plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

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Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

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Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

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Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

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Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

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Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

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Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

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Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

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Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

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Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

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Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

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Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

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Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

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Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

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Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

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van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

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Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

(Francisco and Pinotti 2000) The Lauraceae is betterknown for producing a range of alkaloids (eg Gibbs1974) and as a dominant family in the rainforest flora ofAustralia has been much studied for its toxic and potentiallymedicinal alkaloids (Webb 1949 1952 Collins et al 1990)Of 39 species tested in the Lauraceae family in this surveyonly B collina was cyanogenic (Appendix) Given theapparent rarity of cyanogenesis in the family and eventhe order Laurales it is difficult to speculate as to thebiosynthetic precursor of the cyanogenic constituent inB collina To the authorsrsquo knowledge only the tyrosine-derived cyanogenic glycoside taxiphyllin has been repor-ted in the Calycanthaceae family within the Laurales(Lechtenberg and Nahrstedt 1999) tyrosine-derivedglycosides are also found in Ranunculaceae in the orderRanunculales which is in the same subclass Magnoliidae(sensu Cronquist 1981) as Laurales

Embelia grayi ST Reynolds (Myrsinaceae)

This is the first report of cyanogenesis in the genusEmbeliamdasha genus of approx 130 speciesmdashand amongthe first in the family Myrsinaceae Furthermore Gibbs(1974) considered cyanogenesis to be unknown in theorder Primulales Subsequent to Gibbs (1974) only tworeports of cyanogenesis in Myrsinaceae could be foundmdashfor Rapanea parviflora (Kaplan et al 1983) and forR umbellata which needs confirmation as cyanogenesiswas only detected after 24 h of tissue incubation(Francisco and Pinotti 2000) Overall there are even afew negative test records for the family Embelia grayi isa vine with diameter up to 9 cm endemic to upland andhighland rainforest in north east Queensland Embeliacaulialata and three Rapanea spp also tested here werenot cyanogenic The order Primulales (sensu Cronquist1981) is in the subclass Dilleniidae which also includesthe orders Malvales and Violales Within the Violalescyanogenesis is common in the Achariaceae (includingFlacourtiaceae) Passifloraceae and Turneraceae familieswhich tend to contain cyanogenic glycosides of the cyclo-pentanoid series (Lechtenberg and Nahrstedt 1999)

Passiflora sp (Kuranda BH12896) (Passifloraceae)

Passiflora sp (Kuranda BH12896) was the only speciesin the Passifloraceae tested in this study It is a vine foundin the lowland rainforests of north east Queensland allAustralian members of the Passifloraceae are climbers orsprawling shrubs (Morley and Toelken 1983) Cyanogen-esis is common within the family Passifloraceae (600species two genera worldwide) and the genus Passiflora(eg Rosenthaler 1919 Tjon Sie Fat 1979a Adsersen et al1988 Olafsdottir et al 1989) Several Australian Passifloraspp were reported to be cyanogenic including P aurantia(Petrie 1912 Smith andWhite 1918 Webb 1952 Gardnerand Bennetts 1956) and the endemic P herbertiana Lindl(Petrie 1912) and implicated in poisoning stock (Smithand White 1918) Within Passifloraceae cyanogenesishas also been reported in Adenia spp and Ophiocaulonspp (Rosenthaler 1919 Tjon Sie Fat 1979a) The specificcyanogenic constituents may be taxonomically diagnostic at

the infrafamilial level (eg occurrence of the rare glycosidepassibiflorin Adsersen et al 1993) Cyanogenic Passiflor-aceae typically contain cyanogenic glycosides withcyclopentenoid ring structures (Seigler et al 1982Spencer and Seigler 1985 Nahrstedt 1987 Lechtenbergand Nahrstedt 1999) although phenylalanine-derived glyc-osides (eg prunasin) have been isolated from Passifloraedulis (Spencer and Seigler 1983 Chassagne et al 1996Seigler et al 2002) and valineisoleucine-derived glycos-ides (eg linamarin lotaustralin) from several Passifloraspp in the subgenus Plectostemma (Spencer et al 1986)

Cyanogenesis in the Proteaceae family

Five of the 20 species tested in the Proteaceae familywere found to be cyanogenic C sublimis FMuellO heterophylla LSSm Helicia australasica FMuellH blakei Foreman and H nortoniana (FMBailey)FMBailey The latter species was opportunistically testedas it was found only outside the plots This is the firstformal report of cyanogenesis in the monospecific generaCardwellia and Opisthiolepis while cyanogenesis has pre-viously been reported in the genus Helicia (H robustaGibbs 1974)

Cardwellia sublimis (lsquothe northern silky oakrsquo) is the onlyspecies in the tribe Cardwelliinae (Hoot and Douglas 1998)This canopy species (to 30m) an important timber tree isendemic to north east Queensland being widely distributedthroughout well-developed lowland to highland rainforest(Hyland et al 2003) Cardwellia sublimis was common toall sites in this study

Opisthiolepis heterophylla (lsquothe blush silky oakrsquo) isendemic and confined to north east Queensland from theKirrama Range to Cooktown It grows to 30m in lowlandto highland rainforest but is most common in upland andhighland rainforest on the Atherton Tableland (Hyland et al2003) The flowers of O heterophylla have been found tobe cyanogenic (E E Conn University of California DavisCA USA pers comm)

The genus Helicia (approx 90 species) occurs through-out Asia and the Pacific with nine species found naturallyin Australia (Foreman 1995) Helicia blakei (lsquoBlakersquos silkyoakrsquo) is endemic to north east Queensland occurring as anunderstorey tree in well-developed upland rainforest(Hyland et al 2003) Helicia australasica is a shrub ortree (3ndash20m) widespread throughout northern Australianthrough to Papua New Guinea It occurs as an understoreytree in well-developed rainforest monsoon forest and dryrainforest (Foreman 1995 Hyland et al 2003) The fruit isknown to be eaten by aborigines (Foreman 1995) Helicianortoniana is also an understorey tree (to 20m) endemicto north east Queensland and found in well-developedlowland and highland rainforest (Cooper and Cooper1994 Foreman 1995 Hyland et al 2003) These datasupport the preliminary results of tests on dried herbariumsamples where only some samples of H australasica andH nortoniana tested positive for cyanide (E E ConnUniversity of California Davis CA USA pers comm)Tests of dried herbarium samples of H blakei howevergave only negative results (E E Conn University of

1026 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

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usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

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hn

soniiFM

uell

Rham

naceae

(5)

UB1

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izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

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TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

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VS

mil

ax

acu

lea

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Smilacaceae

(3)

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VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

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VS

mil

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Smilacaceae

(2)

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mil

ax

gly

cip

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Smilacaceae

(2)

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SH

So

lan

um

ma

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raiFM

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Solanaceae

(1)

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lan

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sp

Solanaceae

(2)

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lan

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Dunal

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(1)

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on

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uell)LSSm

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onaceae

(3)

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SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

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era

ma

coo

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(FM

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Monim

iaceae

(4)

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TSte

noca

rpus

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hite

Proteaceae

(1)

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EEConn(perscomm)ndashve

MELU102196

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ua

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Proteaceae

(1)flwrndash

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EEConn(perscomm)ndashve

VS

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an

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nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

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ella

aust

rali

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ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

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ber

var

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rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

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VS

tryc

hn

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min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

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Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

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itto

nsi

iNoot

Symplocaceae

(2)

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TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

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Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

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MELU102289

TS

yno

um

mu

elle

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Meliaceae

(2)

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TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

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Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

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iflo

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(FM

uell)BHyland

Myrtaceae

(3)

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end

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loiu

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Myrtaceae

(4)

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TS

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sta

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ides

(FM

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Myrtaceae

(1)

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Myrtaceae

(3)

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(4)

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ma

nn

ii(FM

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Myrtaceae

(1)

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no

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mu

mCraven

Myrtaceae

(1)

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(3)

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(1)

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(3)

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102249

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m(FM

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Myrtaceae

(3)

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Taber

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Apocynaceae

(1)

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Tra

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14

78

0)

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(4)

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Ta

sma

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Winteraceae

(3)dry

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ernst

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hite

Theaceae

(2)

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no

rdti

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no

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(2)

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Monim

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(6)

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(3)

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(1)

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(3)

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his

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(1)

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Rubiaceae

(1)

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Myrtaceae

(1)

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enti

lag

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Monim

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(1)

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SH

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kiea

sp

Monim

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(2)

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SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

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SH

Wil

kiea

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(DaviesCreek

LJ

Web

b+

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30)

Monim

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(3)

UB1

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SH

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kiea

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elli

i(FM

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(3)

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m(FM

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Xanthophyllaceae

(3)

UL

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TX

ylop

iam

acc

rea

e(FM

uell)LSSm

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(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

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(2)

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Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

California Davis CA USA pers comm) The inconsistentfindings by Conn are probably the result of using driedherbarium material

Cyanogenesis is considered especially common inthe Proteaceae (Swenson et al 1989 Lechtenberg andNahrstedt 1999) known in a range of genera but par-ticularly in Grevillea and Hakea spp In Australia cyano-genic members of the Proteaceae have been implicated instock poisoning (Gardner and Bennetts 1956) Based on thereports of Gibbs (1974) and Tjon Sie Fat (1979a) and thestudy of Swenson et al (1989) who found 44 of 155 pro-teaceous species tested to be cyanogenic cyanogenesisis most widespread in the subfamily Grevilleoideae Forexample cyanogenesis has been reported in the generaStenocarpus Lomatia Helicia Xylomelum TelopeaMacadamia Hicksbeachia Lambertia Grevillea andXylomelum (Petrie 1912 Smith and White 1918 Hurst1942 Gardner and Bennetts 1956 Gibbs 1974 Swensonet al 1989 Lamont 1993 E E Conn University ofCalifornia Davis CA USA pers comm) all of whichare in the Grevilloiedeae (Hoot and Douglas 1998) Con-sistent with this pattern the cyanogenic genera reportedhere are all in the subfamily Grevilleoideae Cardwelliain the subtribe Cardwelliinae (tribe Knightieae) Opisthio-lepis in the subtribe Buckinghamiinae (tribe Embothrieae)and Helicia in the subtribe Heliciinae (tribe Heliceae) Incontrast there are only a few reports of cyanogenesis withinthe Proteoideae subfamily cyanogenesis was only reportedin single species of Conospermum Petrophile and Protea(Gibbs 1974 Swenson et al 1989 E E Conn Universityof California Davis CA USA pers comm)

The cyanogenic constituents which have been identifiedin comparatively few species appear to be biogeneticallyderived from tyrosine Swenson et al (1989) identifiedthe cyanogenic glycosides in eight species leaves andflowers of several Hakea Leucadendron Grevilleaand Macadamia species were found to contain dhurrinand proteacin (see also Plouvier 1964 Young andHamilton 1966) In this regard the identity of the cyano-genic constituent in the monospecific genera in particularwould be interesting

Prunus turneriana (FMBailey) Kalkman (Rosaceae)

Cyanogenesis is widespread within Rosaceae (Hegnauer1990) and has been much studied in the subfamilyPrunoideae in particular as it contains many cyanogeniccultivated species [eg Prunus domestica L (plum)Armeniaca vulgaris Lam (apricot)] Cyanogenic Rosaceae(in particular Prunus spp) are also a common source ofpoisoning in domestic animals (eg Poulton 1983Schuster and James 1988) Prunus turneriana is one ofonly two Prunus species native to Australia and is a latesuccessional canopy tree species in the lowland and uplandrainforests of far north Queensland Australia The fruits ofthis canopy species are known to be toxic yet are also eatenby cassowaries fruit pigeons Herbert river ringtail possumsand musky-rat kangaroos (Cooper and Cooper 1994)The flesh of the fruit was used raw by the Ngadjonjipeoplemdashthe original inhabitants of the rainforests onthe Atherton Tablelands north Queenslandmdashfor treating

toothache while the toxic kernels were processed for astarchy food (Huxley 2003) Despite its common namelsquoAlmond barkrsquo and phytochemical surveys of Australianrainforest species (eg Webb 1949) cyanogenesis in Pturneriana had not previously been reported Cyanogenicglycosides were found to be distributed throughout all tis-sues and consistent with other species in the Rosaceae arebiosynthetically derived from the amino acid phenylalanine(Moslashller and Seigler 1999) Prunasin was identified as themajor cyanogen in leaf stem root and seed tissues ofP turneriana and amygdalin was restricted to the seedWhat was unusual about P turneriana was the presenceof significant amounts of the (R)-prunasin epimer (S)-sambunigrin in leaf stem and seed tissue whereas roottissue contained only prunasin Refer to Miller et al(2004) for the detailed characterization of cyanogenesisin P turneriana

Brombya platynema FMuell (Rutaceae)

This is the first report of cyanogenesis in the genusBrombya which is a genus of 1ndash2 species endemic toAustralia (Hyland et al 2003) Brombya platynema isendemic to north east Queensland where it occurs as anunderstorey tree in well-developed forest (from sea levelto 1100m asl) (Hyland et al 2003) The family (150genera 1500 species) includes many strongly scentedshrubs and trees and rutaceous species are known fortheir terpenoids and alkaloids (Price 1963 Gibbs 1974Everist 1981) Cyanogenesis is rare in Rutaceae and hasonly been reported in Boronia bipinnate Lindl (leavesRosenthaler 1919) Zieria spp (Hurst 1942 Gibbs 1974Fikenscher and Hegnauer 1977) Zanthoxylum fagara(Adsersen et al 1988) and Loureira cochinchinensisMeissa (Gibbs 1974) Even within the order Rutalescyanogenesis is rare with only a few additional definitivereports of cyanogenesis in the Tremandraceae family(Gibbs 1974) The phenylalanine-derived cyanogenic glyc-osides prunasinsambunigrin and zierin have been isolatedfrom leaves of two Zieria spp (Finnemore and Cooper1936 Fikenscher and Hegnauer 1977) The rare meta-hydroxylated cyanogenic glycoside holocalin was recentlyidentified as the principal cyanogen in leaves ofB platynema traces of prunasin and amygdalin werealso detected (Miller et al 2006b) These data suggestthe possibility that species in this family have cyanogenicglycosides biosynthetically derived from the amino acidphenylalanine

Mischocarpus grandissimus (FMuell) Radlk andMischocarpus exangulatus (FMuell) Radlk (Sapindaceae)

Two of the four species of Mischocarpus tested in thisstudy were found to be cyanogenicmdashM grandissimus andM exangulatusmdashrepresenting the first reports of cyanogen-esis for the genus Mischocarpus is a genus of 15 speciesfound in Asia Malesia and Australia nine species occurnaturally in Australia (Hyland et al 2003) Both speciesare endemic to Queensland M grandissimus is restricted tonorth east Queensland while M exangulatus is also foundon the Cape York Peninsula Queensland Mischocarpus

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1027

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

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Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

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Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

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Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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grandissimus occurs as an understorey tree in well-developed lowland and upland rainforest (sea level to750m asl Hyland et al 2003) Similarly M exangulatus(lsquothe red bell mischocarprsquo) is an understorey tree to 15mfound in well-developed lowland and highland rainforest(sea level to 1100m asl Cooper and Cooper 1994Hyland et al 2003) These were the only cyanogenicspecies identified among the 29 species in the Sapindaceaefamily tested in this study (Appendix) Cyanogenesisis known in the Sapindaceae family however thefamily is best known for cyanolipids in the seed oils ofnumerous species (eg Alectryon spp Allophylus sppCardiospermum spp Sapindus spp Paullinia spp andUngnadia speciosa) (Mikolajczak et al 1970 Seigleret al 1971 Gowrikumar et al 1976 Seigler andKawahara 1976) There are only a few reports ofcyanogenesis in Australian indigenous membersof Sapindaceaemdashfor Dodonaea spp (Hurst 1942 Webb1949) and Alectryon spp (Smith and White 1918Finnemore and Cooper 1938)mdashand with the exceptionof Heterodendrum oleifolium Desf [syn Alectryon oleifo-lius (Desf) SReyn] the cyanogenic constituents in thesespecies have not been characterized In addition to cyanol-ipids in the seeds leucine-derived cyanogenic glycosideshave been characterized from vegetative parts of sapin-daceous species (Seigler et al 1974 Hubel andNahrstedt 1975 1979 Nahrstedt and Hubel 1978)

Further findings

Alocasia brisbanensis (Araceae) was also found to becyanogenic consistent with reports of cyanogenesis innumerous congeneric species (Rosenthaler 1919 TjonSie Fat 1979a) However as a herb it was not includedin the analysis The tyrosine-derived cyanogenic glycosidetriglochinin has been isolated from the closely relatedA macrorrhiza (Nahrstedt 1975)

There were several findings which contradicted previousreports for species In addition to negative results forA scholaris noted above Eupomatia laurina (Eupoma-tiaceae) was reported to be lsquodoubtfully cyanogenicrsquo andCananga odorata (Annonaceae) cyanogenic by Gibbs(1974) but neither species was found to be cyanogenicin this study (based on repeated tests of four and two indi-viduals respectively) Similarly reports of cyanogenesis infruit and leaves of the Australian endemic Davidsonrsquos plum(Davidsonia pruriens) (Petrie 1912 Rosenthaler 1929)were not corroborated both mature leaves and fruit testingnegative in this study Gibbs (1974) also only found neg-ative results for leaves of D pruriens In addition negativetest results for cyanogenesis in this study corroborate pre-vious negative reports for Neolitsea dealbata (syn Litseadealbata) Cayratia acris Morinda jasminoides and Sarco-petalum harveyanum (Gibbs 1974 Appendix) A screeningof Australian Proteaceae family conducted primarily usingdry herbarium material was carried out by E E Conn(University of California Davis CA USA pers comm)and included several species tested in this study As notedabove Conn had some inconsistent and possibly thereforeinconclusive results for a number of species which wereconfirmed by fresh sampling in this study One of eight

samples of Musgravea heterophylla gave a positive reactionafter 12 h in Connrsquos survey a species which was not foundto be cyanogenic here Further testing of M heterophyllais warranted As far as could be discerned the majority ofspecies tested in this study had not previously been testeddespite numerous screenings of Australian and Queenslandplants most notably by Webb (1948 1949)

General findings frequency of cyanogenesis

Of 401 species from 87 families tested in the survey18 (45) species from 13 families were cyanogenicThe proportion of cyanogenic species at the sites rangedfrom 47 to 65 values similar to the frequency of cyano-genic species found in a substantial survey of woody speciesin Costa Rican rainforest (Thomsen and Brimer 1997)mdashtheonly previous study to report the frequency of cyanogenesisstandardized with respect to plant size (dbh gt10 cm) andforest area (7 middot 1 ha plots) Overall Thomsen and Brimer(1997) found that 40 (range 21ndash57 for plots) of401 species from 68 families were cyanogenic and thatcyanogenic stems (dbh gt 5 cm) accounted for 3 oftotal basal area (range 16ndash51) Here the overall propor-tion of total basal area in cyanogenic stems was 73 andranged from 12 to 134 The highest proportions were inhighland rainforest on basalt soil (134) and in lowlandrainforest (116) Highland rainforest on rhyolite had thelowest proportion

Overall at a community level there are few studies withwhich to compare frequencies of cyanogenesis reportedhere for tropical rainforest in north east Queensland Inthe first instance there have been few studies in tropicalsystems but also the screening methodology variesdepending on the research question being addressed be ittaxonomic (eg examining chemical differences in relationto proposed phylogenetic relationships) or ecological(eg the role of secondary compounds in plantndashanimal inter-actions) For example while the survey of Thomsen andBrimer (1997) was standardized with respect to plant sizeand forest area the few surveys in other tropical systemshave focused on plantndashanimal interactions and have notscreened in a standardized fashion Only 23 (one species)was found to be cyanogenic in the screening of gt90 of theflora (n = 43 species) in a species-poor seasonal cloud forestin India (Mali and Borges 2003) In contrast in a surveyexamining the frequency of cyanogenesis in relation toenvironmental factors and insect density along a transectfrom the shoreline to an inland lagoon in a neotropicalwoodland (lsquorestingarsquo) Kaplan et al (1983) found 25 species(23) of 108 species screened to be cyanogenic They alsoreported variable test results for a further 49 species (for n =2ndash16 individuals) elevating the proportion of cyanogenicspecies to 68 a value which requires further examinationbefore interpretation as the sampling strategy (eg plantsize random sampling strategy life form and transect area)was unclear and some uncertainty with regard to picratepaper test results was expressed by the authors (Kaplanet al 1983)

Adsersen et al (1988) compared frequencies of cyano-genesis within the endemic and non-endemic flora of theGalapagos Islands two floras subject to different suites of

1028 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

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Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

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TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

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Myrtaceae

(4)

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yzyg

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Myrtaceae

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john

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Myrtaceae

(3)

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mo

no

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mCraven

Myrtaceae

(1)

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BHyland

Myrtaceae

(3)

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MELU102159

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yzyg

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m(CTW

hite)

BHyland

Myrtaceae

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yzyg

ium

wes

aBHyland

Myrtaceae

(3)

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MELU102096

102249

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wil

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Taber

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Apocynaceae

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14

78

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Winteraceae

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(6)

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zNon-w

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oftaxonomic

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Ryp

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quantitativelyforthepresence

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aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

herbivores over a period of time They screened fresh andherbarium specimens of a significant proportion (65) ofthe flora from the Archipelago and reported 81 ofendemic species and 53 of native speciesmdashthosewhich also occur on the South American mainlandmdashtobe cyanogenic Interestingly they also reported a further22 of native species and 30 of endemic species torelease cyanide in the presence of a crude mix of b-glycosidases (including 5 b-glucuronidase from snails)suggesting that a large number of species contain cyano-genic glycosides but lack the catabolic b-glycoside enzymeThis contrasts with the findings of several other studieswhere the addition of non-specific b-glycosidases orpectinase during qualitative testing did not alter the fre-quency of positive results (eg Petrie 1912 Thomsenand Brimer 1997 Buhrmester et al 2000 Lewis andZona 2000 but see Conn et al 1985) Similarly in thisstudy all samples were spontaneously cyanogenic withoutthe addition of pectinase from Rhizopus spp indicatingthat non-cyanogenic individuals probably lacked both thecyanogenic glycoside and b-glycosidase or possibly thatpectinase was not able to catalyse the cyanogenesis in thesespecies It is perhaps noteworthy that the greater frequencyof cyanogenesis reported by Conn et al (1985) in responseto the addition of b-glycosidase (emulsin) was in a surveysolely of the genus Acacia indicating that such a responsemay vary among taxa

It is important to note that the frequencies reported inall of these surveys were apparently based on the testing ofsingle specimens of the vast majority of species (Adsersenet al 1988 Thomsen and Brimer 1997 Mali and Borges2003) Similarly while the present study aimed to test atleast three individuals of each species in duplicate (ie withand without added enzyme) only one individual of manyspecies was encountered (Appendix) Furthermore mostspecies here were also tested in both wet and dry seasonsA range of studies report variable positive and negative testresults among sample sizes as small as n = 2 (eg Kaplanet al 1983 Thomsen and Brimer 1997) Given such poly-morphism for cyanogenesis (see also Aikman et al 1996)the reported frequencies in these surveys may underestimateoverall the actual proportion of cyanogenic species in plantcommunities The reported frequency of cyanogenesis mayalso vary with the plant part tested In the Costa Rican studyThomsen and Brimer (1997) reported a greater frequency ofcyanogenesis among reproductive plant parts than leavesas did Buhrmester et al (2000) in populations of Sambucuscanadensis (elderberry) in Illinois Consistent with thattrend individuals of species with weakly cyanogenic leavesincluding some which produced negative FA paper resultshad higher concentrations of glycosides in flowers or fruitshowever overall only a small number of reproductivetissues were tested so limited comparison can be drawn(Table 1)

The frequency of cyanogenesis varies betweentaxonomic groups and with life form Cyanogenesis is con-sidered especially common in some plant families (egRosaceae Euphorbiaceae Passifloraceae and ProteaceaeLechtenberg and Nahrstedt 1999) and rare or absent inothers (eg Lauraceae and Araliaceae Gibbs 1974

Hegnauer 1989) a trend which may in part reflect differ-ential intensity of testing among taxonomic groups In thisstudy the frequency of cyanogenesis among the dominantplant families varied In the Proteaceae family five of20 (25) species were cyanogenic while two of 29(69) species in the Sapindaceae and one of 39 (25)species in the Lauraceae were cyanogenic (Table 1) In ascreening of Australian Acacia 69 of 360 species werecyanogenic (Conn et al 1985) Cyanogenesis appears to berare among palms only two species (12) of 155 speciesof palms (108 genera) were found to be cyanogenic(Lewis and Zona 2000) No cyanogenic palm specieswere identified in this study

Concentrations of cyanogenic glycosides

Several of the 18 cyanogenic species detected in thisstudy contained concentrations of cyanogenic glycosidesamong the highest reported for leaves of woody speciesMost notably tree species E sericopetalus C grayi and Pturneriana had foliar concentrations of cyanogenic glycos-ides up to 52 49 and 48mgCNgndash1 d wt respectively(Table 1) Similarly Webber (1999) recorded concentra-tions up to 5mgCN gndash1 d wt in the tree species R javanicaindividuals of that species occurring within the surveyarea of this study had a lower mean concentration of18mgCN gndash1 d wt (Table 1) These high concentrationsare substantially greater than the majority of values reportedfor foliage from a range of tropical and temperate taxa Forexample concentrations up to 11mgHCNgndash1 d wt werereported in the tropical shrub Turnera ulmifolia (Shoreand Obrist 1992 Schappert and Shore 1999) while thehighest concentrations in naturally occurring populations ofAustralian Eucalyptus spp were 259mgCNgndash1 d wt and316mgCNgndash1 d wt for E cladocalyx and E yarraensisrespectively (Gleadow and Woodrow 2000a Goodgerand Woodrow 2002) Foliar concentrations of between166 and 378mgHCNg1 d wt have been recorded incultivated Prunus spp (Santamour 1998) To our know-ledge possibly the highest foliar cyanogenic glycosideconcentration among naturally occurring woody specieswas reported in the tropical proteaceous species Panopsiscostaricensis by Thomsen and Brimer (1997) who meas-ured 2150mgHCNkgndash1 f wt (approximately equivalent to72mgHCNg1 d wt using a conversion based on the meanfoliar water content of several species in this study whichwas 70) The age of the leaves analysed was not specifiedhowever and it should also be noted that this value wasdetermined using picrate papers and reflectrometry a lessdependable method more sensitive to the presence ofinterfering substances (Brinker and Seigler 1989)

Assessing the ecological significance of the rangeof cyanogenic glycoside concentrations recorded here isdifficult While the high concentrations in several species(eg gt2mgCNg1 d wt Table 1) would almost certainlyconstitute toxic levels important in defencemdashplants withgt600 mgHCNg1 d wt are considered potentially danger-ous to livestock for example (Haskins et al 1987)mdashtheecological significance of lower concentrations (eg approx8ndash50mgCNg1 d wt) is harder to determine This reflects

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1029

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the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

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Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

Dow

nloaded from httpsacadem

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TS

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theFlacourtiaceae

(Chase

eta

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zNon-w

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anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

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nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

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sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

the fact that despite the well-documented effectiveness ofcyanogenesis in defence against generalist herbivores(eg Jones 1998 Gleadow and Woodrow 2002) overallthere is no known particular concentration at whichcyanogenic compounds are effective in herbivore deter-rence This is in part because unfortunately many herbivorystudies only report the presence or absence of cyanogenesisand not the actual concentrations of cyanogenic glyc-osides Moreover the efficacy of cyanogenesis as adefence depends not only on the concentration of cyano-genic glycosides but also on the physiology morphologyand behaviour of the consumer (Gleadow and Woodrow2002)

Intra-plant variation in cyanogenic glycoside content

In tropical forests it is estimated that up to 70 of a leafrsquoslifetime damage occurs while expanding (Coley and Barone1996) This differential intensity of herbivory among oldand young leaves and the frequent observation that defencecompounds tend to be concentrated in plant tissues of highervalue to reproduction or growth (eg young leaves) is sum-marized in the Optimal Allocation Theory (OAT) ofdefence This theory predicts that the most vulnerableand valuable plant partsmdashthose susceptible to attack andmost likely to contribute to growth and reproductive fitnesssuch as reproductive structures and young leavesmdashwill bemore defended (McKey 1974 Rhoades 1979)

Results here add to the already substantial body of workon mostly temperate cyanogenic species consistent withthe predictions of the OAT (eg Martin et al 1938Dement and Mooney 1974 Cooper-Driver et al 1977Shore and Obrist 1992 Dahler et al 1995 Thomsenand Brimer 1997 Gleadow et al 1998 Gleadow andWoodrow 2000b) In all species where young leaveswere sampled they contained significantly higher concen-trations of cyanogenic glycosides than old leaves Thistrend was most apparent in species that had low cyanogencontent in old leaves (eg C sublimis C myrianthusO heterophylla and P australiana) (Table 1) Moreoverin some cases individuals of these species appeared only toinvest in cyanogenic glycoside defence in young leavesacyanogenic old leaves were seemingly reliant more onphysical toughness Thus leaf age is an important consid-eration when assigning the cyanogenic phenotype

Again consistent with the OAT reproductive tissuessuch as floral buds flowers and fruitsseeds tended tohave high total cyanogen content (Table 1) This patternhas been commonly reported among cyanogenic species(eg Spencer and Seigler 1983 Selmar et al 1991Selmar 1993b Thomsen and Brimer 1997 Webber1999) One notable exception to this was the low to negli-gible concentrations of cyanogenic glycosides in matureseeds of C sublimis unlike the fleshy seeds of many rain-forest species C sublimis seeds are dry and papery Theabsence of cyanogenesis in mature seeds of proteaceousGrevillea spp was reported by Lamont (1993) in specieswith cyanogenic foliage and flowers The higher concentra-tions in floral tissues of C sublimis is consistent with pre-vious reports for proteaceous species which tend to have

high concentrations of cyanogenic glycosides in flowerswhile leaves may have low total cyanogen content or beacyanogenic (eg Smith and White 1918 Tjon Sie Fat1979a Lamont 1993)

Intra-population variation in cyanogenic glycoside content

All individuals of the majority of species were cyano-genic albeit with low concentrations of cyanogenicglycosides in some instances Negative results with FApapers for old leaves were only obtained for individualsof three of the 18 cyanogenic species In the populationof one of these species B platynema 50 of individualswere determined to be acyanogenic with cyanogenic gly-coside concentrations much less than the 8mgCN g1 d wtthreshold The two other exceptions were C myrianthusand P australiana Unlike B platynema cyanogenesis inindividuals of these species varied qualitatively with leafage and plant part Thus assigning the acyanogenic pheno-type in these species was problematic

This developmental trend towards differences inexpression of cyanogenic potential has been reported pre-viously cyanogenesis is known be affected by plant agegrowth phase as well as the plant part used (Jones 1972Gibbs 1974 Seigler 1991) Consequently as noted earlierstudies have reported a greater frequency of cyanogenesiswhen testing reproductive tissues young foliage and shootscompared with old leaves (eg Gibbs 1974 Aikman et al1996 Thomsen and Brimer 1997 Buhrmester et al 2000Mali and Borges 2003) These findings emphasize theimportance of only comparing leaves of a similar agewhen classifying individuals according to the presence orabsence of cyanogenesis

Aside from the three species mentioned above noacyanogenic individuals were identified in populations ofother cyanogenic species While the small sample sizes formost species reduced the likelihood of encountering anacyanogenic individual even within populations of themore abundant species such as B collina (n = 46 allsites) C sublimis (n = 31 at all sites) and R javanica(n = 249 Webber 1999) no acyanogenic individualswere detected In the latter example the quantitative screen-ing of gt800 individuals of R javanica failed to detect anacyanogenic individual in several distinct populations(Webber 2005) This number (800) was substantiallygreater than the number of individuals predicted byGleadow and Woodrow (2000a) and Goodger et al(2002) (n = 95ndash100) that would need to be sampled tocapture an acyanogenic individual assuming a similar gen-etic system for cyanogenesis to Trifolium repens (Hugheset al 1988) and an estimate of the rarity of a species (orpolymorph) (McArdle 1990) The floristic heterogeneity ofthe rainforest makes sampling large populations challen-ging it is noteworthy however that others have detectedpolymorphism for cyanogenesis in tropical studies basedon very small sample sizes (eg n = 2) (Kaplan et al1983 Thomsen and Brimer 1997) These acyanogenicmorphs determined only by indicator paper tests in thesestudies were not verified by quantitative assay as forB platynema here

1030 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

Islands ecological and evolutionary aspects OIKOS 66 511ndash520Adsersen A Adsersen H Brimer L 1988 Cyanogenic constituents in

plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

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TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

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Theaceae

(2)

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Rubiaceae

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Sapindaceae

(3)

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Sapindaceae

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Proteaceae

(3)

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Rubiaceae

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1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Conclusions

In summary the findings of this survey indicate thatcyanogenesis is an important yet little studied chemicaldefence in tropical rainforests The identification of specificcyanogens in but a few of the cyanogenic species first repor-ted here has yielded novel findings Given the large numberof new reports for species belonging to plant familiesor orders in which cyanogenesis has been little reportedthe ongoing characterization of cyanogenic constituents inthese species will potentially be of both phytochemical andchemotaxonomic significance In addition preliminary dataon intra-population variation in cyanogenesis here suggestthat ontogenetic variation in cyanogenesis and polymorph-ism for cyanogenesis merit further investigation in tropicalrainforest species

ACKNOWLEDGEMENTS

The authors thank Professor Eric Conn (University ofCalifornia Davis) for sharing unpublished data on theProteaceae family Nicole Middleton and volunteers atthe University of Melbourne Herbarium for mountingand databasing specimens Andrew Ford (CSIRO TropicalForest Research Centre Atherton Queensland) for assist-ance with species name changes and Wendy Cooper andSteve McKenna for additional taxonomic assistance duringfield work We also thank Dr John Kanowski (GriffithUniversity Queensland) and Andrew Graham (TFRCAtherton) for advice in the selection of field sites andMarisa Collins Bruce Webber Ross Waller JenniferFox Gavin Smith and Joel Youl for assistance in thefield We thank the Australian Canopy Crane ResearchFacility for access to forest on the site Field work wassupported by an Individual Award from the QueenrsquosTrust for Young Australians to REM and was conductedunder the Queensland Department of Environment andHeritage Scientific Purposes Permit number F100027099SAA and Department of Natural Resources Permitsto Collect numbers 1542 1716 1409 and 1663

LITERATURE CITEDAdsersen A Adsersen H 1993 Cyanogenic plants in the Galapagos

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plants from the Galapagos Islands Biochemical Systematics andEcology 16 65ndash77

Aikman K Bergman D Ebinger J Seigler D 1996 Variation ofcyanogenesis in some plant species of the Midwestern United StatesBiochemical Systematics and Ecology 24 637ndash645

Brimer L Cicalini AR Federici F Petruccioli M 1995Beta-glycosidaseas a side activity in commerical pectinase preparations of fungalorigin The hydrolysis of cyanogenic glycosides Italian Journal ofFood Sciences 4 387ndash394

Brinker AM Seigler DS 1989 Methods for the detection and quantitativedetermination of cyanide in plant materials Phytochemical Bulletin21 24ndash31

Buhrmester RA Ebinger JE Seigler DS 2000 Sambunigrin andcyanogenic variability in populations of Sambucus canadensisL CaprifoliaceaeBiochemical Systematics and Ecology 28 689ndash695

Cantino PD 1992 Evidence for a polyphyletic origin of the LabiataeAnnals of the Missouri Botanical Garden 79 361ndash379

CFSAN Centre for Food Safety and Applied Nutrition 2003 USDepartment of Health and Human Services Poisonous Plant Databasehttpwwwcfsanfdagovdjwplantoxhtml Accessed 16 October2003

Chase MW Zmarzty S Lledo MD Wurdack KJ Swensen SMFay MF 2002 When in doubt put it in Flacourtiaceae a molecularphylogenetic analysis based on plastid rbcL DNA sequences KewBulletin 57 141ndash181

Chassagne D Crouzet J Bayonove CL Baumes RL 1996 Identificationand quantification of passion fruit cyanogenic glycosides Journal ofAgricultural and Food Chemistry 44 3817ndash3820

Coley PD 1983Herbivory and defensive characteristics of tree species in alowland tropical rainforest Ecological Monographs 53 209ndash233

Coley PD 1988Effects of plant growth rate and leaf lifetime on the amountand type of anti-herbivore defense Oecologia 74 531ndash536

Coley PD Barone JA 1996 Herbivory and plant defenses in tropicalforests Annual Review of Ecology and Systematics 27 305ndash335

Coley PD Kursar TA 1996 Anti-herbivore defenses of young tropicalleaves physiological constraints and ecological trade-offs InMulkey SS Chazdon RL Smith AP eds Tropical forest plant eco-physiology New York Chapman and Hall 305ndash336

Coley PD Bryant JP Chappin FS III 1985 Resource availability andplant antiherbivore defense Science 230 895ndash899

Collins DJ Culvenor CCJ Lamberton JA Loder JW Price JR 1990Plants for medicines a chemical and pharmacological survey ofplants in the Australian region Melbourne CSIRO Publications

Conn EE 1981Cyanogenic glycosides In Conn EE ed The biochemistryof plants New York Academic Press Inc 479ndash499

Conn EE 1991 The metabolism of a natural product lessons learnedfrom cyanogenic glycosides Planta Medica 57 S1ndashS9

Conn EE Maslin BR Curry S Conn ME 1985 Cyanogenesis inAustralian species of Acacia Survey of herbarium specimens andliving plantsWestern Australian Herbarium Research Notes 10 1ndash13

Cooper W Cooper WT 1994 Fruits of the rain forest A guide to fruitsin Australian tropical rain forests Surry Hills NSW AustraliaRD Press

Cooper-Driver GA Finch S Swain T 1977 Seasonal variation in sec-ondary plant compounds in relation to the palatability of Pteridiumaquilinum Biochemical Systematics and Ecology 5 177ndash183

Cronquist A 1981 An integrated system of classification of floweringplants New York Columbia University Press

Dahler JM McConchie CA Turnbull CGN 1995 Quantification ofcyanogenic glycosides in seedlings of three Macadamia Proteaceaespecies Australian Journal of Botany 43 619ndash628

Davies AG Bennett EL Waterman PG 1988 Food selection by twoSouth-east Asian colobine monkeys Preshytis rubicunda andPresbytis melalophos in relation to plant chemistryBiological Journalof the Linnean Society 34 33ndash56

Dement WA Mooney HA 1974 Seasonal variation in the productionof tannins and cyanogenic glycosides in the Chaparral shrubHeteromeles arbutifolia Oecologia 15 65ndash76

Dickenmann R 1982 Cyanogenesis in Ranunculus montanus sl from theSwiss Alps Bericht des Geobotanischen Institutes ETH 49 56ndash75

Everist SL 1981 Poisonous plants of Australia Sydney Australia Angusand Robertson

Eyjolfsson R 1971 Constitution and stereochemistry of lucumin acyanogenic glycoside from Lucuma mammosa Gaertn Acta ChemicaScandanavia 25 1898ndash1900

Feeny P 1976 Plant apparency and chemical defense In Wallace JWMansell RL eds Biochemical interaction between plants andinsects New York Plenum Press 1ndash40

Feigl F Anger V 1966 Replacement of benzidine by copper ethylaceto-acetate and tetra base as spot-test reagent for hydrogen cyanide andcyanogen Analyst 91 282ndash284

Fikenscher LH Hegnauer R 1977 Cyanogenese bei den Cormophyten12 Mitteilung Chaptalia nutans eine stark cyanogene PlanzeBrasiliens Planta Medica 31 266ndash269

Finnemore H Cooper JM 1936 Cyanogenetic glucosides in Australianplants Part 4mdashZieria laevigata Journal of the Royal Society ofNew South Wales 70 175ndash182

FinnemoreH CooperJM 1938Thecyanogenic constituentsofAustralianand other plants Journal of the Society of Chemistry and Industry 57167ndash169

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1031

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

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Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

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Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

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Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

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Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

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Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

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Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

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Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

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Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

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Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

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Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

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Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

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Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

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Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

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Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

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Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

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Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Finnemore H Cox CB 1928Cyanogenetic glucosides inAustralianplantsJournal of the Royal Society of New South Wales 62 369ndash378

Finnemore H Cox CB 1929 Cyanogenic glucosides in Australian plantsPart II A Eremophila maculata the native fuchsia Journal of theRoyal Society of New South Wales 63 172ndash178

Foreman DB 1995 Helicia Flora of Australia 16 393ndash399Forster PI Williams JB 1996 Apocynaceae Flora of Australia 28

104ndash196Francisco IA Pinotti MHP 2000 Cyanogenic glycosides in

plants Brazilian Archives of Biology and Technology 43 487ndash492Gardner CA Bennetts HW 1956 The toxic plants of Western Australia

Perth Australia West Australian Newpapers LtdGartlan JS McKeyDB Waterman PG Mbi CN StruhsakerTT 1980A

comparative study of the phytochemistry of two African rain forestsBiochemical Systematics and Ecology 8 401ndash422

Gibbs RD 1974 Chemotaxonomy of flowering plants Vol 1ndash3 Montrealand London Queenrsquos University Press

Gleadow RM Woodrow IE 2000aPolymorphism in cyanogenicglycosidecontent andcyanogenicb-glucosidaseactivity innatural populationsofEucalyptus cladocalyx Australian Journal of Plant Physiology 27693ndash699

Gleadow RM Woodrow IE 2000b Temporal and spatial variation incyanogenic glycosides in Eucalyptus cladocalyx Tree Physiology20 591ndash598

Gleadow RM Woodrow IE 2002 Constraints on effectiveness ofcyanogenic glycosides in herbivore defence Journal of ChemicalEcology 28 1297ndash1309

Gleadow RM Foley WJ Woodrow IE 1998 Enhanced CO2 alters therelationship between photosynthesis and defence in cyanogenicEucalyptus cladocalyx F Muell Plant Cell and Environment 2112ndash22

Goodger JQD Woodrow IE 2002 Cyanogenic polymorphism as anindicator of genetic diversity in the rare species Eucalyptus yarraensisMyrtaceae Functional Plant Biology 29 1445ndash1452

Goodger JQD Capon RJ Woodrow IE 2002 Cyanogenic poly-morphism in Eucalyptus polyanthemos Schauer subsp vestita LJohnson and K Hill Myrtaceae Biochemical Systematics andEcology 30 617ndash630

Gowrikumar G Mani VVS Lakshminarayana G 1976 Cyanolipids inSapinus emarginatus seed oil Phytochemistry 15 1566ndash1567

Graham AW Hopkins MS Maggs J 1995 Succession and disturbance inthe remnant rainforest type complex notophyll vine forest on basalttype 5b I Vegetation map and explanatory notesAtherton AustraliaCSIRO Division of Wildlife and Ecology Tropical Research CentreVM11295-6

Hadi S Bremmer JB 2001 Initial studies on alkaloids from Lombokmedicinal plants Molecules 6 117ndash129

Hall N Wainwright RW Wolf LJ 1981 Summary ofmeteorological data in Australia no 6 CSIRO Australian Divisionof Forest Research

Hartley TG Dunstone EA Fitzgerald JS Johns SR Lamberton JA1973 A survey of New Guinea plants for alkaloids Lloydia 36217ndash319

Hartmann T Witte I Ehmke A Theuring C Rowell-Rahier MPasteels JM 1997 Selective sequestration and metabolism of plantderived pyrrolizidine alkaloids by chrysomelid leaf beetles Phyto-chemistry 45 489ndash497

Haskins FA Gorz HJ Johnson BE 1987 Seasonal variation in leafhydrocyanic acid potential of low- and high-dhurrin Sorghums CropScience 27 903ndash906

Hegnauer R 1966 Chemotaxonomie der pflanzen Eine Ubersicht uberdie Verbreitung und die systematische Bedeutung der PflanzenstoffeVol 4 Basel Birkhauser Verlag

Hegnauer R 1973 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 6 Basel Birkhauser Verlag

Hegnauer R 1986 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 7 Basel Birkhauser Verlag

Hegnauer R 1989 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 8 Basel Birkhauser Verlag

Hegnauer R 1990 Chemotaxonomie der pflanzen Eine Ubersicht uber dieVerbreitung und die systematische Bedeutung der PflanzenstoffeVol 9 Basel Birkhauser Verlag

Hickel A Hasslacher M Griengl H 1996Hydroxynitrile lyases functionsand properties Physiologia Plantarum 98 891ndash898

Hoot SB Douglas AW 1998 Phylogeny of the Proteaceae based on atpBand atpBndashrbcL intergenic spacer region sequences AustralianSytematic Botany 11 301ndash320

Hubel W Nahrstedt A 1975 Ein neues cyanglykosid aus Heterodendronoleaefolium Phytochemistry 14 2723ndash2725

Hubel W Nahrstedt A 1979 Cardiosperminsulfatemdasha sulfur containingcyanogenic glucoside from Cardiospermum grandiflorum Tetrahed-ron Letters 45 4395ndash4396

Hughes MA 1991 The cyanogenic polymorphism in Trifolium repensL white clover Heredity 66 105ndash115

Hughes MA Sharif AL Alison-Dunn M Oxtoby E 1988 Themolecularbiology of cyanogenesis In Evered D Harnett S eds Cyanidecompounds in biology Chichester John Wiley amp Sons 111ndash130

Hurst E 1942 The poison plants of New South Wales Sydney AustraliaPoison Plants Committee NSW

Huxley M 2003 Ngadjon names and uses of some rainforest plants httpwwwkooriusydeduaungadjonjiFoodtable1html Accessed 16February 2004

Hyland BPM Whiffin T Christophel DC Gray B Elick RW 2003Australian rain forest plants trees shrubs vines CollingwoodCSIRO Publishing Australia

Janzen DH Waterman PG 1984 A seasonal census of phenolics fibreand alkaloids in foliage of forest trees in Costa Rica some factorsinfluencing their distribution and relation to host selection bySphingidae and SaturniidaeBiological Journal of the Linnean Society21 439ndash454

Janzen DH Doerner ST Conn EE 1980 Seasonal constancy of intra-population variation of HCN content of Costa Rican Acaciafarnesiana foliage Phytochemistry 19 2022ndash2023

Jaroszewski JW Olafsdottir ES 1987 Monohydroxylated cyclopenten-one cyanohydrin glucosides of Flacourtiaceae Phytochemistry 263348ndash3349

Jensen SR Nielsen BJ 1973Cyanogenic glucosides in Sambucus nigraLActa Chemica Scandanavia 27 2661ndash2662

Jensen SR Nielsen BJ 1986 Gynocardin in Ceratiosicyos laevisAchariaceae Phytochemistry 25 2349ndash2350

Jones DA 1972Cyanogenic glycosides and their function InHarborne JBed Phytochemical ecology London Academic Press 103ndash124

Jones DA 1998Why are somany food plants cyanogenicPhytochemistry47 155ndash162

Kaplan MAC Figueiredo MR Gottlieb OR 1983 Variation incyanogenesis in plants with season and insect pressure BiochemicalSystematics and Ecology 11 367ndash370

Kojima M Poulton JE Thayer SS Conn EE 1979 Tissue distributionsof dhurrin and of enzymes involved in its metabolism in leaves ofSorghum bicolor Plant Physiology 63 1022ndash1028

Kursar TA Coley PD 2003 Convergence in defense syndromes of youngleaves in tropical rainforests Biochemical Systematics and Ecology31 929ndash949

Lamont BB 1993 Injury-induced cyanogenesis in vegetative andreproductive parts of two Grevillea species and their F1 hybridAnnals of Botany 71 537ndash542

Lechtenberg M Nahrstedt A 1999Cyanogenic glycosides In IkanR edNaturally occurring glycosides Chichester UK John Wiley amp Sons147ndash191

Levin DA 1976 Alkaloid-bearing plants an ecogeographic perspectiveAmerican Naturalist 110 261ndash284

Levin DA York BM Jr 1978 The toxicity of plant alkaloids anecogeographic perspective Biochemical Systematics and Ecology 661ndash76

Lewis CE Zona S 2000 A survey of cyanogenesis in palms ArecaceaeBiochemical Systematics and Ecology 28 219ndash228

Mabberley DJ 1990 The plant book Cambridge UK CambridgeUniversity Press

Mali S Borges RM 2003 Phenolics fibre alkaloids saponins andcyanogenic glycosides in a seasonal cloud forest in IndiaBiochemicalSystematics and Ecology 31 1221ndash1246

1032 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Martin JH Couch JF Briese RR 1938 Hydrocyanic acid content ofdifferent parts of the sorghum plant Journal of the American Societyof Agronomy 30 725ndash734

Maslin BR Dunn JE Conn EE 1988 Cyanogenesis in Australian speciesof Acacia Phytochemistry 27 421ndash428

McArdle BH 1990 When are rare species not there OIKOS 57 276ndash277McKey D 1974 Adaptive patterns in alkaloid physiology American

Naturalist 108 305ndash320McKey D Waterman PG Gartlan JS Struhsaker TT 1978 Phenolic

content of vegetation in two African rain forests ecologicalimplications Science 202 61ndash64

Mikolajczak KL Smith CR Jr Tjarks LW 1970 Cyanolipids ofCardiospermum halicacabum L and other Sapindaceous seed oilsLipids 5 812ndash817

Miller RE 2004 Cyanogenesis in Australian tropical rainforests resourceallocation to a nitrogen-based defence PhD Thesis The University ofMelbourne Australia

Miller RE Gleadow RM Woodrow IE 2004 Cyanogenesis in tropicalPrunus turneriana characterisation variation and response to lowlight Functional Plant Biology 31 491ndash503

Miller RE McConville MJ Woodrow IE 2006a Cyanogenic glycosidesfrom the rare Australian endemic rainforest tree Clerodendrum grayi(Lamiaceae) Phytochemistry 67 43ndash51

Miller RE Simon J Woodrow IE 2006b Cyanogenesis in the Australiantropical rainforest endemic Brombya platynema (Rutaceae) chemicalcharacterisation and polymorphism Functional Plant Biology (inpress)

Moslashller BL Seigler DS 1999 Biosynthesis of cyanogenic glycosidescyanolipids and related compounds In Singh KK ed Plant aminoacids biochemistry and biotechnology New York Marcel Dekker563ndash609

Morley BD Toelken HRE 1983 Flowering plants in AustraliaAdelaide Australia Rigby Publishers

Munir AA 1989 A taxonomic revision of the genus Clerodendrum LVerbenaceae in Australia Journal of the Adelaide Botanic Gardens11 101ndash173

Nahrstedt A 1975 Verteilung von prunasin in fruchtstielen vonPrunus avium L und der einfluszlig von IES auf dengesamtgehalt an prunasin Biochemie und Physiologie der Pflanzen167 597ndash600

Nahrstedt A 1980 Absence of cyanogenesis from DroseraceaePhytochemistry 19 2757ndash2758

Nahrstedt A 1987 Recent developments in the chemistry distribution andbiology of the cyanogenic glycosides In Hostettmann K Lea PJ edsBiologically active natural products Oxford Clarendon Press213ndash234

Nahrstedt A Hubel W 1978 Die cyanogenen verbindungen vonHeterodendron oleaefolium Phytochemistry 17 314ndash315

Nix HA 1991Biogeography pattern and process InNixHA SwitzerMAKowari I eds Rainforest animals Atlas of vertebrates endemic toAustraliarsquos wet tropics Canberra Australian National Parks andWildlife Service 11ndash40

Olafsdottir ES Andersen JV Jaroszewski JW 1989 Cyanohydringlycosides of Passifloraceae Phytochemistry 28 127ndash132

Osunkoya OO Ash JE Graham AW Hopkins MS 1993 Growth oftree seedlings in tropical rain forests of North Queensland AustraliaJournal of Tropical Ecology 9 1ndash18

Pammel LH 1911 A manual of poisonous plants Chiefly of easternNorth America with brief notes on economic and medicinal plantsand numerous illustrations Cedar Rapids Iowa USA The TorchPress

Patton CA Ranney TG Burton JD Walgenbach JF 1997 Naturalpest resistance of Prunus taxa to feeding by adult Japanese beetlesrole of endogenous allelochemicals in host plant resistance Journalof the American Society for Horticultural Science 122 668ndash672

Petrie JM 1912 Hydrocyanic acid in plants Part I Its distribution in theAustralian flora Proceedings of the Linnean Society of New SouthWales 37 220ndash234

Petrie JM 1913 Hydrocyanic acid in plants I Proceedings of the LinneanSociety of New South Wales 37 220ndash234

Petrie JM 1914 Hydrocyanic acid in plants II Proceedings of the LinneanSociety of New South Wales 38 624ndash638

Petrie JM 1920 Cyanogenesis in plants IV The hydrocyanic acid ofHeterodendronmdasha fodder plant of New South Wales Proceedingsof the Linnean Society of New South Wales 45 447ndash459

Plouvier V 1964 Investigation of polyalcohols and cyanogenicheterosides in certain Proteaceae Compte Rendu Hebdomadaire desSeances de LrsquoAcademie des Sciences 259 665ndash666

Poulton JE 1983 Cyanogenic compounds in plants and their toxic effectsIn Keeler RF Tu WT eds Plant and fungal toxins New YorkDekker 117ndash157

Poulton JE 1990 Cyanogenesis in plants Plant Physiology 94 401ndash405Poulton JE Li CP 1994 Tissue level compartmentation of R-amygdalin

and amygdalin hydrolase prevents large-scale cyanogenesis inundamaged Prunus seeds Plant Physiology 104 29ndash35

Price JR 1963 The distribution of alkaloids in the Rutaceae In Swain Ted Chemical plant taxonomy London Academic Press 429ndash452

Rhoades DF 1979 Evolution of plant chemical defense againstherbivores In Rosenthal GA Janzen DH eds Herbivores theirinteraction with secondary plant metabolites New York AcademicPress 3ndash54

Rosenthaler L 1919 Die Verbreitung der Blausaure im PflanzenreichJournal Suisse de Pharmacie 57 279ndash283 307ndash313 324ndash329341ndash346

Rosenthaler L 1929 Beitrage zur Blausaure-Frage Neue Blausaure-Pflanzen Pharmaceutica Acta Helvetiae 4 196ndash199

Santamour Jr FS 1998 Amygdalin in Prunus leaves Phytochemistry 471537ndash1538

Saupe SG Seigler DS Escalante-Semerena J 1982 Bacterialcontamination as a cause of spurious cyanide tests Phytochemistry21 2111ndash2112

Schappert PJ Shore JS 1999 Effects of cyanogenesis polymorphism inTurnera ulmifolia on Euptoieta hegesia and potential Anolispredators Journal of Chemical Ecology 25

Schuster JL James LF 1988 Some other major poisonous plants ofthe Western United States In James LF Ralphs MH Darwin DBeds The ecology and economic impact of poisonous plants onlivestock production London Westview Press 295ndash307

Seigler DS 1976a Plants of Oklahoma and Texas capable of producingcyanogenic compounds Proceedings of the Oklahoma Academy ofScience 56 95ndash100

Seigler DS 1976b Plants of the Northeastern United States that producecyanogenic compounds Economic Botany 30 395ndash407

Seigler DS 1991 Cyanide and cyanogenic glycosides In Rosenthal GABerenbaum MR eds Herbivores their interactions with secondaryplant metabolites Academic Press San Diego 35ndash77

Seigler DS 1994 Phytochemistry and systematics of the EuphorbiaceaeAnnals of the Missouri Botanical Garden 81 380ndash401

Seigler DS Kawahara W 1976 New reports of cyanolipids fromSapindaceous plants Biochemical Systematics and Ecology 4263ndash265

Seigler DS Seaman F Mabry TJ 1971 New cyanogenetic lipids fromUngnadia speciosa Phytochemistry 10 485ndash487

Seigler DS Eggerding C Butterfield C 1974 A new cyanogenicglycoside from Cardiospemum hirsutum Phytochemistry 132330ndash2332

Seigler DS Coussio JD Rondina RVD 1979 Cyanogenic plants fromArgentina Journal of Natural Products 42 179ndash182

Seigler DS Spencer KC Statler WS Conn EE Dunn JE 1982TetraphyllinB and epitetraphyllinB sulphates novel cyanogenic gluc-osides fromPassiflora caerulea andP alato-caeruleaPhytochemistry21 2277ndash2285

Seigler DS Pauli GF Nahrstedt A Leen R 2002 Cyanogenicallosides and glucosides from Passiflora edulis and Carica papayaPhytochemistry 60 873ndash882

Selmar D 1993a Apoplastic occurrence of cyanogenic b-glucosidases andconsequences for themetabolismofcyanogenicglucosides InEsenAed The biochemistry and molecular biology of b-glucosidasesAmerican Chemical Society Washington DC 191ndash204

Selmar D 1993b Transport of cyanogenic glucosides linustatin uptake byHevea cotyledons Planta 191 191ndash199

Selmar D Lieberei R Junqueira N Biehl B 1991 Changes incyanogenic glucoside content in seeds and seedlings ofHevea speciesPhytochemistry 30 2135ndash3140

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1033

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

Shore JS Obrist CM 1992 Variation in cyanogenesis within andamong populations and species of Turnera series CanaligeraeTurneraceae Biochemical Systematics and Ecology 20 9ndash16

Smith F White CT 1918 An interim census of cyanophoric plants in theQueensland flora Proceedings of the Royal Society of Queensland 3084ndash90

Smolenski SJ Silinis H Farnsworth NR 1974 Akaloid screening IIILloydia 36 359

Spencer KC Seigler DS 1983 Cyanogenesis of Passiflora edulis Journalof Agricultural and Food Chemistry 31 794ndash796

Spencer KC Seigler DS 1985 Cyanogenic glycosides and the sys-tematics of the Flacourtiaceae Biochemical Systematics and Ecology13 421ndash431

Spencer KC Seigler DS Nahrstedt A 1986 Linamarin lotaustralinlinustatin and neolinustatin from Passiflora species Phytochemistry25 645ndash647

Swanholm CE St John H Scheuer PJ 1960 A survey for alkaloids inHawaiian plants II Pacific Science 14 68ndash74

Swenson WK Dunn JE Conn EE 1989 Cyanogenesis in the ProteaceaePhytochemistry 28 821ndash823

Taskova R Mitova M Evstatieva L Ancev M Peev D Handjieva NBankova V Popov S 1997 Iridoids flavonoids and terpenoids astaxonomic markers in Lamiaceae Scrophulariaceae and RubiaceaeBocconea 5 631ndash636

Thomsen K Brimer L 1997 Cyanogenic constituents in woody plantsin natural lowland rain forest in Costa Rica Biological Journal ofthe Linnean Society 121 273ndash291

Tjon Sie Fat L 1978 Contribution to the knowledge of cyanogenesis inAngiosperms 2 Communicatioin Cyanogenesis in CampanulaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 81 126ndash131

Tjon Sie Fat L 1979a Contribution to the knowledge of cyanogenesis inangiosperms Cyanogene Verbindingen bij Poaceae CommelinaceaeRanunculaceae en Campanulaceae PhD Thesis Leiden UniversityThe Netherlands

Tjon Sie Fat L 1979b Contribution to the knowledge of cyanogenesis inAngiosperms 14 Communication Cyanogenesis in RanunculaceaeProceedings of the Koninklijke Nederlandse Akadamie VanWetenschappen Series C 82 197ndash209

Tracey JC 1982 The vegetation of the humid tropical region of NorthQueensland Melbourne Australia CSIRO

Tracey JG Webb LJ 1975 Vegetation of the humid tropical region ofNorth Queensland Indooroopilly Queensland Rainforest EcologyUnit CSIRO Australia Division of Plant Industry

Urbanska K 1982 Polymorphism of cyanogenesis in Lotus alpinus inSwitzerland I Small-scale variability in phenotypic frequencies uponacidic silicate and carbonate Bericht des Geobotanischen InstitutesETH 49 35ndash55

van Valen F 1978 Contribution to the knowledge of cyanogenesis inangiosperms Planta Medica 34 408ndash413

van Wyk B-E Whitehead CS 1990 The chemotaxonomic significanceof prunasin in Buchenroedera Fabaceae-Crotalarieae South AfricanJournal of Botany 56 68ndash70

Wagstaff SJ Olmstead RG 1997Phylogeny of Labiatae andVerbenaceaeinferred from rbcL sequences Systematic Botany 22 165ndash179

Wagstaff SJ Hickerson L Spangler R Reeves PA Olmstead RG1998 Phylogeny in Labiatae s l inferred from cpDNA sequencesPlant Systematics and Evolution 209 265ndash274

Wajant H Riedel D Benz S Mundry K-W 1994 Immunocytologicallocalization of hydroxynitrile lyases from Sorghum bicolor L andLinum usitatissimum L Plant Science 103 145ndash154

Waterman PG Ross AM Jr Bennett EL Davies AG 1988 Acomparison of the floristics and leaf chemistry of the tree flora intwo Malaysian rain forests and the influence of leaf chemistry onpopulations of colobinemonkeys in theOldWorldBiological Journalof the Linnean Society 34 1ndash32

Webb LJ 1948 Guide to the medicinal and poisonous plants ofQueensland CSIRO Bulletin 232 1ndash202

Webb LJ 1949 An Australian phytochemical survey I Alkaloids andcyanogenetic compounds in Queensland plants CSIRO Bulletin241 1ndash56

Webb LJ 1952 An Australian phytochemical survey II Alkaloids inQueensland flowering plants CSIRO Bulletin 268 1ndash99

Webb LJ Tracey JG 1981 The rainforests of northern AustraliaIn Groves RH ed Australian vegetation Cambridge CambridgeUniversity Press 67ndash101

Webb LJ Tracey JG Williams WT 1972 Regeneration andpattern in the sub-tropical rainforest Journal of Ecology 60675ndash95

Webber BL 1999 Cyanogenesis in the tropical rainforest tree Ryparosajavanica Honours Thesis The University of Melbourne

Webber BL 2005 Plantndashanimal interactions and plant defence in therainforest tree Ryparosa PhD Thesis The University of MelbourneAustralia

Webber BL Woodrow IE 2004Cassowary frugivory seed defleshing andfruit fly infestation influence the transition from seed to seedling inthe rare Australian rainforest tree Ryparosa sp nov 1 AchariaceaeFunctional Plant Biology 31 505ndash516

Young RL Hamilton RA 1966 A bitter principle in Macadamia nutsIn Proceedings of the 6th annual meeting Hawaii MacadamiaProducers Association 27ndash30

Zheng L Poulton JE 1995 Temporal and spatial expression ofamygdalin hydrolase and R-+-mandelonitrile lygase in black cherryseeds Plant Physiology 109 31ndash39

APPENDIX

Summary of all species tested for cyanogenesis within6 middot 200m2 plots at five sites in uplandhighland (U) orlowland (L) tropical rainforest on sites contrasting insoil type basalt sites at Lamins Hill (B1) and LonglandsGap (B2) and sites on granite at Mt Nomico (G) on rhyoliteat Longlands Gap (R) and on metamorphic substrate nearCape Tribulation (M) Species are listed in alphabeticalorder Life forms tree (T) shrub (SH) herb (H) vine (V)treefern (TF) palm (P) and hemi-epiphyte (HE) The results(+ or ndash) for tests using FeiglndashAnger papers and approx1 g f wt leaf tissue with and without the addition ofpectinase (+ndash enz) for n individuals of each species arelisted All tests were carried out using most recently fullyexpanded leaves and in most instances also using youngleaves In some cases leaf tips (tips) only a few fruit(ft) or flowers (flwr) were tested Previous findings forspecies and in some cases genera are noted (eg Proteaceaespecies tested by E E Conn University of CaliforniaDavis CA USA pers comm based on herbarium speci-mens) Species included in phytochemical screenings ofQueensland rainforest taxa (alkaloids CN Webb 1948Webb 1949) are noted most were not tested for CN(DNT) Canopy species or rare species for which no samplewas obtained were included in floristic analysis but werenot tested for cyanogenesis (DNT) Lodgement numbers atBrisbane (BRI) and The University of Melbourne (MELU)herbaria are given

1034 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TA

caci

ace

lsaTindale

Mim

osaceae

(3)

UR

Webb(1949)DNT

TA

cmen

agra

veole

ns(FM

Bailey)LSSm

Myrtaceae

(3)

LM

MELU102318

TA

cmen

are

saBHyland

Myrtaceae

(2)

UB2R

TA

cro

nyc

hia

aci

du

laFM

uell

Rutaceae

(2)

UG

Webb(1949)DNT

TA

cro

nyc

hia

acr

onyc

hio

ides

(FM

uell)TGHartley

Rutaceae

(3)

UG

TA

cro

nyc

hia

cra

ssip

eta

laTGHartley

Rutaceae

(1)

UB2

SH

Acr

onyc

hia

pa

rvifl

ora

CTW

hite

Rutaceae

(5)

UB1R

SH

Act

eph

ilasp(W

ooroonooranNP

PI

Fors

ter+

PIF

171

51)

Euphorbiaceae

(5)

UB1

TA

den

an

ther

ap

avo

nin

aL

Mim

osaceae

(1)

LM

TA

ga

this

atr

op

urp

ure

aBHyland

Araucariaceae

(3)

UR

TA

ga

this

mic

rost

ach

yaJFBaileyamp

CTW

hite

Araucariaceae

(1)

UG

TA

ga

this

rob

ust

a(CM

oore

exFM

uell)FM

Bailey

Araucariaceae

(1)

UB1

SH

Ag

laia

mer

idio

na

lisCM

Pannell

Meliaceae

(13)

UL

B1M

MELU102332

TA

gla

iasa

pin

din

a(FM

uell)Harms

Meliaceae

(2)

LM

TA

gla

iato

men

tosa

Teijsmamp

Binn

Meliaceae

(5)

UL

B1B2GM

MELU102275

TA

lan

giu

mvi

llo

sum

subsp

po

lyo

smo

ides

(FM

uell)Bloem

b

Alangiaceae

(3)

UB1

Webb(1949)DNT

MELU102329

TA

llo

xylo

nfl

am

meu

mPHW

estonamp

Crisp

Proteaceae

(2)

UB2

TA

llo

xylo

nw

ickh

am

ii(W

Hillex

FM

uell)

PHW

estonamp

Crisp

Proteaceae

(4)

UGB2

HA

loca

sia

bri

sba

nen

sis(FM

Bailey)Domin

zAraceae

++

UB1

Hurst(1942)

TA

lph

ito

nia

wh

itei

Braid

Rham

naceae

(3)

UG

MELU102103

HA

lpin

iaa

rcti

flora

(FM

uell)Benth

Zingiberaceae

(1)

UB1

HA

lpin

iam

od

esta

FM

uellex

KSchum

Zingiberaceae

(1)

UB2

TA

lsto

nia

sch

ola

ris(L)RBr

Apocynaceae

(10)

LM

Gibbs(1974)reported

+CNWebb(1949)DNT

BRI578809

MELU102127

102257

SH

Aly

xia

ilic

ifoli

asubsp

il

icif

oli

aFM

uell

Apocynaceae

(3)

UG

VA

lyxi

asp

ica

taRBr

Apocynaceae

(2)

UB1R

Webb(1949)DNT

TA

nti

des

ma

ero

stre

FM

uellex

Benth

Euphorbiaceae

(6)

UL

B1GRM

MELU102113

SH

An

tirh

easp(M

tMissery

LW

Jes

sup+

GJD

31

36)

Rubiaceae

(3)

UB2

TA

nti

rhea

ten

uifl

ora

FM

uellex

Benth

Rubiaceae

(4)

UL

B1GM

TA

po

dyt

esb

rach

ysty

lisFM

uell

Icacinaceae

(6)

UB1GR

MELU102188

TA

rch

iden

dro

nra

mifl

oru

m(FM

uell)Kosterm

Mim

osaceae

(4)

UL

B1M

TA

rch

iden

dro

nva

illa

nti

i(FM

uell)FM

uell

Mim

osaceae

(3)

UB2G

MELU102186

TA

rch

iden

dro

nw

hit

eiICNielsen

Mim

osaceae

(1)

UB1

SH

Ard

isia

bif

ari

aCTW

hiteamp

WDFrancis

Myrsinaceae

(2)

UB1

SH

Ard

isia

bre

viped

ata

FM

uell

Myrsinaceae

(7)frtndash

UL

B1B2GRM

MELU102148

TA

rgyr

od

endro

nsp(Boonjee

BH

21

39R

FK)

Sterculiaceae

(9)

UB1

TA

ryte

rap

au

cifl

ora

STReynolds

Sapindaceae

(4)

UL

B1M

SH

Atr

act

oca

rpus

hir

tus(FM

uell)Puttock

Rubiaceae

(7)

UL

B1M

Webb(1949)DNT

SH

Atr

act

oca

rpus

mer

ikin

(FM

Bailey)Puttock

Rubiaceae

(3)

UB1B2

TA

ura

nti

carp

ap

ap

yra

ceaLCayzer

Crisp

ampITelford

Pittosporaceae

(2)

UB2G

VA

ust

rob

ail

eya

sca

nden

sCTW

hite

Austrobaileyaceae

(6)flwrndash

UB1

SH

Au

stro

ma

ttha

eael

ega

nsLSSm

Monim

iaceae

(3)

UGR

TA

ust

rom

uel

lera

trin

ervi

aCTW

hite

Proteaceae

(4)

UL

B1M

VA

ust

rost

eenis

iast

ipula

ris(CTW

hite)

Jessup

Fabaceae

(4)

UB1G

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1035

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TB

ala

no

ps

au

stra

lia

naFM

uell

Balanopaceae

(3)

URG

Webb(1949)DNT

MELU102216

102108

TB

eils

chm

iedia

bancr

oft

ii(FM

Bailey)CTW

hite

Lauraceae

(3)

UL

B1GM

Webb(1949)DNT

MELU102219

TB

eils

chm

iedia

cast

risi

nen

sisBHyland

Lauraceae

(2)

LM

TB

eils

chm

iedia

coll

inaBHyland

Lauraceae

++

UB1B2GR

BRI578804

MELU102299

102300

TB

eils

chm

iedia

oli

gandra

LSSm

Lauraceae

(3)

UG

TB

eils

chm

iedia

recu

rvaBHyland

Lauraceae

(4)

UB1G

TB

eils

chm

iedia

toora

m(FM

Bailey)BHyland

Lauraceae

(10)

UB1B2G

MELU102211

TB

ob

eam

yrto

ides

(FM

uell)Valeton

Rubiaceae

(8)

UGR

MELU102105102110

SH

Bo

wen

iasp

ecta

bil

isHookex

Hookf

Stangeriaceae

(6)

UL

B1M

TB

rach

ych

ito

na

ceri

foli

us(A

Cunn

exGDon)Macarthur

Sterculiaceae

(3)

UL

B2M

TB

rack

enri

dg

eaa

ust

rali

an

aFM

uell

Ochnaceae

(4)

UL

GRM

MELU102322

SHT

Bre

ynia

cern

ua(Poir)MuellArg

Euphorbiaceae

(2)

UB2G

Webb(1949)1948

B

ob

lon

gif

oli

aMuell

Arg+C

NT

Bro

mbya

pla

tyn

emaFM

uell

Rutaceae

++

LM

Webb(1949)DNT

BRI578818

MELU102283

102313

TB

ubbia

sem

ecarp

oid

es(FM

uell)BLBurtt

Winteraceae

(5)

UB1G

V(P)

Cala

mu

sa

ust

rali

sMart

Arecaceae

(1)

UB1

VC

ala

mu

sm

otiFM

Bailey

Arecaceae

(1)

UB1

TC

ald

clu

via

au

stra

lien

sis(Schltr)Hoogland

Cunoniaceae

(1)

UB1

TC

alo

ph

yllu

mco

sta

tum

FM

Bailey

Clusiaceae

(3)

UB2R

TC

an

an

ga

od

ora

ta(Lam

)Hookfamp

Thomson

Annonaceae

(2)

LM

TC

an

ari

um

mu

elle

riFM

Bailey

Burseraceae

(1)

UG

MELU102250

TC

an

thiu

msp

Rubiaceae

(1)

UG

TC

ara

llia

bra

chia

ta(Lour)Merr

Rhizophoraceae

(1)

LM

TC

ard

wel

lia

sub

lim

isFM

uell

Proteaceae

++

UL

B1B2GRM

EEConn(perscomm)

1of7+v

eBRI578806

MELU102280102281

TC

arn

arv

on

iaa

rali

ifo

liavar

m

on

tan

aBHyland

Proteaceae

(3)

UB2GR

EEConn

(perscomm)ndashve

MELU102282

VC

arr

on

iap

rote

nsa

(FM

uell)Diels

Menispermaceae

(5)

UL

B1B2M

MELU102213

TC

ase

ari

aco

stula

taLW

Jessup

Salicaceaedagger

(4)

UB1B2GR

Webb(1949)DNT

TC

ase

ari

ad

all

ach

iiFM

uell

Salicaceaedagger

(1)

LM

TC

ase

ari

ag

rayi

LW

Jessup

Salicaceaedagger

(2)dry

UB2

TC

ast

an

osp

erm

um

au

stra

leACunnamp

CFraserex

Hook

Fabaceae

(3)

LM

Webb(1949)DNT

TC

ast

an

osp

ora

alp

ha

nd

ii(FM

uell)FM

uell

Sapindaceae

(3)

UB1B2

MELU102130

VC

ayr

ati

asa

pon

ari

a(Seemamp

Benth)Domin

Vitaceae

(1)

LM

C

acr

isGibbs(1974)ndashCN

TC

elti

sp

an

icu

lata

(Endl)Planch

Ulm

aceae

(1)

LM

VC

epha

lara

lia

ceph

alo

botr

ys(FM

uell)Harms

Araliaceae

(2)

UB1B2

TC

erato

pet

alu

msu

ccir

ub

rum

CTW

hite

Cunoniaceae

(4)

UB2G

Webb(1949)DNT

TC

erber

afl

ori

bu

nd

aKSchum

Apocynaceae

(1)

UL

GM

Webb(1949)DNT

TC

hio

na

nth

us

axi

lla

risRBr

Oleaceae

(4)

UB1G

MELU102327

TC

inn

am

om

um

lau

ba

tiiFM

uell

Lauraceae

(3)

UB1GR

Webb(1949)DNT

MELU102210

VC

issu

sh

ypo

gla

uca

AGray

Vitaceae

(2)

UG

VC

issu

sp

enn

iner

vis(FM

uell)Planch

Vitaceae

(1)

UL

RM

1036 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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VC

issu

sst

ercu

liif

oli

a(FM

uellex

Benth)Planch

Vitaceae

(4)

UB1B2G

MELU102325

VC

issu

svi

nosa

Jackes

Vitaceae

(3)

UB1B2G

MELU102195

TC

itro

nel

lasm

yth

ii(FM

uell)RAHoward

Icacinaceae

(4)

UL

B1GM

MELU102143

TC

lao

xylo

nte

ner

ifo

liu

m(Baill)FM

uell

Euphorbiaceae

(1)

UB1

TC

leis

tan

thu

sm

yria

nth

us(H

assk)Kurz

Euphorbiaceae

++

LM

BRI578811

MELU102122

102123102258

TC

lero

den

dru

mgra

yiMunir

Verbenaceae

++

UB1B2G

Other

Cle

rod

end

rum

spp+C

N(G

ibbs1974

TjonSie

Fat1979a

Adsersen

eta

l1988)

BRI578815

MELU102115

102116102117

TC

nes

mo

carp

on

da

syan

tha(Radlk)Adem

aSapindaceae

(2)

UG

VC

on

na

rus

con

choca

rpu

sFM

uell

Connaraceae

(1)

LM

SH

Cord

ylin

ep

etio

lari

s(D

omin)Pedley

Laxmanniaceae

(2)

UB1

SH

Cord

ylin

esp

Laxmanniaceae

(1)

UG

TC

ory

noca

rpus

crib

bia

nus(FM

Bailey)LSSm

Corynocarpaceae

(1)

UB1

C

laev

iga

ta+C

N(W

ebb1948)

SH

Cri

spil

ob

ad

isp

erm

a(SM

oore)Steenis

Alseuosm

iaceae

(4)

UG

TC

rypto

cary

aa

ng

ula

taCTW

hite

Lauraceae

(4)

UB1GR

MELU102102

TC

rypto

cary

aco

coso

ides

BHyland

Lauraceae

(3)

UB2G

MELU102157102268

TC

rypto

cary

aco

rru

ga

taCTW

hiteamp

WDFrancis

Lauraceae

(3)

UB1B2GR

MELU102147

TC

rypto

cary

ad

ensi

flo

raBlume

Lauraceae

(2)

UGR

MELU102104102269

TC

rypto

cary

ag

ran

disBHyland

Lauraceae

(3)

UL

B1B2GM

MELU102247

TC

rypto

cary

ah

ypo

spo

dia

FM

uell

Lauraceae

(1)

LM

TC

rypto

cary

ala

evig

ata

Blume

Lauraceae

(2)

UL

GM

TC

rypto

cary

ale

uco

ph

ylla

BHyland

Lauraceae

(1)

UB2R

MELU102248

TC

rypto

cary

ali

vid

ula

BHyland

Lauraceae

(2)

UGR

MELU102095

TC

rypto

cary

am

ack

inn

on

ianaFM

uell

Lauraceae

(5)

UL

B1GM

MELU102212

TC

rypto

cary

am

ela

no

carp

aBHyland

Lauraceae

(5)

UB1B2

TC

rypto

cary

am

urr

ayi

FM

uell

Lauraceae

(2)

UL

B1M

TC

rypto

cary

ao

bla

taFM

Bailey

Lauraceae

(2)

UL

B1M

Webb(1949)DNT

MELU102099

TC

rypto

cary

ap

leu

rosp

erm

aCTW

hiteamp

WDFrancis

Lauraceae

(4)

UB1G

Webb(1949)DNT

MELU102270

TC

rypto

cary

ap

uti

daBHyland

Lauraceae

(4)

UGR

MELU102144

TC

rypto

cary

asa

cchara

taBHyland

Lauraceae

(3)

UB2R

TC

rypto

cary

asm

ara

gd

inaBHyland

Lauraceae

(3)

UB2R

MELU102193

TC

up

an

iopsi

sfl

ag

elli

form

is(FM

Bailey)

Radlkvar

fl

ag

elli

form

isSapindaceae

(4)

UB1

MELU102129

TC

up

an

iopsi

ssp

Sapindaceae

(1)

LM

TF

Cya

thea

reb

ecca

e(FM

uell)Domin

Cyatheaceae

(2)

UGR

TC

yclo

ph

yllu

mm

ult

iflo

rum

STReynoldsamp

RJFHend

Rubiaceae

(1)

UB1

Canth

ium

vacc

inif

oli

um

FMuell+C

NWebb

(1949)1948

TD

ap

hn

an

dra

rep

and

ula

(FM

uell)FM

uell

Monim

iaceae

(2)

UB1B2G

TD

arl

ingia

da

rlin

gia

na(FM

uell)LASJohnson

Proteaceae

(2)

UG

EEConn(perscomm)ndashve

TD

arl

ingia

ferr

ug

inea

JFBailey

Proteaceae

(3)ndash

UB1B2

EEConn(perscomm)ndashve

TD

avi

dso

nia

pru

rien

sFM

uell

Davidsoniaceae

(2)

UG

Webb(1949)DNT+C

N(Rosenthaler1929Hurst

1942)ndashCN

Gibbs(1974)

MELU102151102152

TD

ecasp

erm

um

humile(G

Don)AJScott

Myrtaceae

(1)dry

LM

TD

ela

rbre

am

ich

iea

na(FM

uell)FM

uell

Araliaceae

(4)

UB1G

VD

end

rotr

op

he

vari

an

s(Blume)

Miq

Santalaceae

(1)

UR

VD

erri

ssp(D

aintree

DE

B

oyl

an

d+4

69)

Fabaceae

(1)dry

LM

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1037

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APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nued

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

VD

esm

os

goez

eanus(FM

uell)Jessup

Annonaceae

(2)

UG

VD

ich

ap

eta

lum

pa

pua

nu

m(Becc)Boerl

Dichapetalaceae

(4)

UB1

MELU102326

TD

ino

sper

ma

stip

ita

tum

(CTW

hiteamp

WDFrancis)

TGHartley

Rutaceae

(3)

UB1

TD

iosp

yro

scu

pu

losa

FM

uell

Ebenaceae

(1)

LM

TD

iosp

yro

sh

ebec

arp

aACunnex

Benth

Ebenaceae

(1)dry

UB1

MELU102307

TD

iplo

glo

ttis

ber

nie

an

aSTReynolds

Sapindaceae

(2)

LM

TD

iplo

glo

ttis

bra

ctea

taLeenh

Sapindaceae

(1)

UG

TD

ory

pho

raa

rom

ati

ca(FM

Bailey)LSSm

Monim

iaceae

(5)

UL

B1B2M

Webb(1949)DNT

TD

ysoxy

lum

all

iace

um

Blume(Blume)

Meliaceae

(1)

LM

TD

ysoxy

lum

arb

ore

scen

s(Blume)

Miq

Meliaceae

(2)

UL

B1M

TD

ysoxy

lum

klander

iFM

uell

Meliaceae

(3)

UB1B2R

TD

ysoxy

lum

opposi

tifo

lium

FM

uell

Meliaceae

(3)

UB2G

MELU102273

TE

laeo

carp

us

ba

ncr

oft

iiFM

uellamp

FM

Bailey

Elaeocarpaceae

(3)

LM

TE

laeo

carp

us

eum

un

diFM

Bailey

Elaeocarpaceae

(1)

UGR

MELU102125

TE

laeo

carp

us

fove

ola

tusFM

uell

Elaeocarpaceae

(2)

UB1R

TE

laeo

carp

us

gra

ha

miiFM

uell

Elaeocarpaceae

(1)

LM

TE

laeo

carp

us

larg

iflo

ren

sCTW

hitesubsp

larg

iflo

ren

sElaeocarpaceae

(4)

UB1G

TE

laeo

carp

us

rum

ina

tusFM

uell

Elaeocarpaceae

(3)

UB2G

TE

laeo

carp

us

seri

cop

eta

lusFM

uell

Elaeocarpaceae

++

UGR

BRI578817

MELU102135

102304

TE

laeo

carp

ussp(M

tBellenden

Ker

LJB

18

336)

Elaeocarpaceae

(3)

UB2GR

MELU102304

VE

mb

elia

cau

lia

lata

STReynolds

Myrsinaceae

(1)dry

LM

VE

mb

elia

gra

yiSTReynolds

Myrsinaceae

++

UB1B2

BRI578803

MELU102267

TE

nd

iand

rab

essa

ph

ilaBHyland

Lauraceae

(2)

UB1

TE

nd

iand

rad

iels

ian

aTeschn

Lauraceae

(2)

UG

TE

nd

iand

rah

ypo

tep

hra

FM

uell

Lauraceae

DNT

LM

TE

nd

iand

rale

pto

den

dro

nBHyland

Lauraceae

(7)

UL

B1M

MELU102293

TE

nd

iand

ram

icro

neu

raCTW

hite

Lauraceae

(3)

LM

TE

nd

iand

ram

on

oth

yraBHylandsubsp

m

on

oth

yra

Lauraceae

(3)

UB1B2

TE

nd

iand

ram

on

tan

aCTW

hite

Lauraceae

DNT

UB2

TE

ndia

ndra

palm

erst

onii(FM

Bailey)

CTW

hiteamp

WDFrancis

Lauraceae

(2)

UB2G

Webb(1949)DNT

MELU102294

TE

nd

iand

rasa

nke

yanaFM

Bailey

Lauraceae

(3)

UB1B2

MELU102155

TE

nd

iand

rasi

der

oxy

lonBHyland

Lauraceae

(3)

UB2

MELU102109

TE

nd

iand

raw

olf

eiBHyland

Lauraceae

(3)shtndash

UB1G

MELU102295

TE

ndosp

erm

um

myr

mec

ophil

um

LSSm

Euphorbiaceae

(1)dry

L

MV

En

tad

ap

ha

seo

loid

es(L)Merr

Mim

osaceae

(1)

LM

HE

Epip

rem

num

pin

natu

m(L)Engl

Araceae

(2)

LM

VE

ryci

be

cocc

inea

(FM

Bailey)Hoogl

Convolvulaceae

(3)

LM

TE

ryth

roxy

lum

ecari

na

tum

Burckex

Hoch

Erythroxylaceae

(2)

UB1

Webb(1949)DNT

MELU102225

SH

Eu

po

ma

tia

ba

rba

taJessup

Eupomatiaceae

(2)dry

UL

B1M

SHT

Eu

po

ma

tia

lauri

naRBr

Eupomatiaceae

(4)

UB2G

Gibbs(1974)lsquodoubtfully

cyanogenicrsquoWebb

(1949)DNT

MELU102331

VE

ust

rep

hu

sla

tifo

liu

sRBrex

Ker

Gaw

lPhilesiaceae

(2)

UL

B1M

TF

ag

raea

cam

ba

gei

Domin

Gentianaceae

(4)

LM

MELU102317

1038 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

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TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TF

ag

raea

fagra

eace

a(FM

uell)Druce

Gentianaceae

(3)

UB1B2GR

MELU102323

TF

icus

conges

taRoxb

Moraceae

(1)

LM

TF

icu

scr

ass

ipes

FM

Bailey

Moraceae

(1)

UB1

TF

icu

sd

estr

uen

sFM

uellex

CTW

hite

Moraceae

(1)

UB2G

TF

icu

sle

pto

cla

daBenth

Moraceae

DNT

UB1B2

MELU102107

VF

icu

sp

an

ton

ian

aKing

Moraceae

(1)

LM

TF

icu

sp

leuro

carp

aFM

uell

Moraceae

(1)

UB1

TF

icu

str

ira

dia

taCorner

Moraceae

(1)

LM

VF

lag

ella

ria

ind

icaL

Flagellariaceae

++

UL

B1GM

+CN

(1912)

BRI578816

MELU102259

102296

TF

lin

der

sia

bo

urj

oti

an

aFM

uell

Rutaceae

(4)

UL

B2GRM

Webb(1949)DNT

TF

lin

der

sia

bra

yley

an

aFM

uell

Rutaceae

(1)

UB2

Webb(1949)DNT

TF

lin

der

sia

laev

ica

rpaCTW

hiteamp

WDFrancis

Rutaceae

(2)

UG

MELU102222

TF

lin

der

sia

pim

ente

lia

naFM

uell

Rutaceae

(2)

UGR

Webb(1949)DNT

MELU102132

102106

TF

on

tain

eap

icro

sper

maCTW

hite

Euphorbiaceae

(2)

UB2

TF

ranci

scoden

dro

nla

uri

foli

um

(FM

uell)BHylandamp

Steenis

Sterculiaceae

(10)

UB1G

VF

reyc

inet

iaex

cels

aFM

uell

Pandanaceae

(3)

UB1B2

VF

reyc

inet

iasc

anden

sGaudich

Pandanaceae

(1)

UB1

TG

alb

uli

mim

ab

acc

ata

FM

Bailey

Him

antandraceae

(3)

UB1B2GR

Webb(1949)DNT

fruitsndashCN

(Bailey1909

inWebb1948)

TG

arc

inia

gib

bsi

aeSM

oore

Clusiaceae

(7)

UB1

MELU102126

TG

arc

inia

warr

eniiFM

uell

Clusiaceae

(4)

LM

TG

ard

enia

ovu

lari

sFM

Bailey

Rubiaceae

(5)

UL

GM

Webb(1949)DNT

MELU102319

TG

evu

ina

ble

asd

ale

i(FM

uell)Sleumer

Proteaceae

(3)

UB1G

EEConn(perscomm)ndashve

MELU102292

TG

illb

eea

ad

enop

eta

laFM

uell

Cunoniaceae

(3)

UB1G

TG

loch

idio

nh

arv

eya

nu

mDomin

var

h

arv

eya

nu

mEuphorbiaceae

(1)

UG

Webb(1949)DNT

TG

loch

idio

nh

yla

nd

iiAiryShaw

Euphorbiaceae

(1)

UB2G

Webb(1949)DNT

TG

mel

ina

fasc

icu

liflo

raBenth

Lam

iaceae

(2)

UL

GM

Webb(1949)DNT

MELU102261

102262102263

TG

om

ph

an

dra

au

stra

lian

aFM

uell

Icacinaceae

(2)

LM

MELU102333

TG

on

ioth

ala

mu

sa

ust

rali

sJessup

Annonaceae

(6)

UB1

MELU102185

102290

TG

oss

iad

all

ach

iana(FM

uell)NSnow

ampGuymer

Myrtaceae

(5)

UB1B2G

MELU102220

TG

oss

iam

yrsi

no

carp

a(FM

uell)

NSnow

ampGuymer

Myrtaceae

(1)

UB2

TG

oss

iag

rayi

iNSnow

ampGuymer

Myrtaceae

(2)

UG

MELU102146

TG

revi

llea

ba

iley

an

aMcG

ill

Proteaceae

(1)

LM

Webb(1952)Hurst(1942)

Gre

vill

easpp+C

N

EEConn(perscomm)ndashve

TG

uio

aa

cuti

foli

aRadlk

Sapindaceae

(2)

UL

B2M

TG

uio

ala

sio

neu

raRadlk

Sapindaceae

(2)

UB1G

TG

uio

am

on

tan

aCTW

hite

Sapindaceae

(2)

UR

MELU102150

SH

Gym

no

sta

chys

an

cep

sRBr

Araceae

(2)

UL

B2M

VG

yno

chto

des

sp(Lam

bRange

JW

398)

Rubiaceae

(2)

UGR

TH

alf

ord

iake

nd

ack

(Montrouz)Guillaumin

Rutaceae

(3)

UB1B2GR

Webb(1949)DNT

MELU102112

SH

Ha

plo

stic

ha

nth

ussp(CoopersCreek

B

Gra

y2

43

3)

Annonaceae

(7)

LM

MELU102291

SH

Ha

plo

stic

ha

nth

ussp(Topaz

LW

Jes

sup

52

0)

Annonaceae

(7)

UB1

MELU102328

SH

Ha

rpull

iafr

ute

scen

sFM

Bailey

Sapindaceae

(1)

UB2

SHT

Ha

rpull

iarh

ytic

arp

aCTW

hiteamp

WDFrancis

Sapindaceae

(10)

UL

B1GM

Webb(1949)DNT

TH

edyc

ary

alo

xoca

rya(Benth)WDFrancis

Monim

iaceae

(4)

UG

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1039

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APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

TH

elic

iaa

ust

rala

sica

FM

uell

Proteaceae

++

LM

EEConn(perscomm)

flwr+v

eH

ro

bust

a+C

NPam

mel1911in

Webb(1949)

1948Gibbs(1974)

BRI578805

MELU102284

TH

elic

iab

lake

iForeman

Proteaceae

++

UB1

EEConn(perscomm)ndashve

BRI578807

MELU102285

TH

elic

iala

min

gto

nia

na(FM

Bailey)

CTW

hiteex

LSSm

Proteaceae

(3)

UB2

EEConn(perscomm)ndashve

MELU102214

TH

ern

and

iaa

lbifl

ora

(CTW

hite)

Kubitzki

Hernandiaceae

(4)

LM

Webb(1949)DNT

MELU102314

VH

ibb

erti

asc

an

den

s(W

illd)Dryand

Dilleniaceae

(1)

UR

VH

ipp

ocr

ate

ab

arb

ata

FM

uell

Celastraceae

(4)

UL

B1M

VH

oya

sp

Asclepiadaceae

(1)

LM

TH

yla

nd

iad

ock

rill

iiAiryShaw

Euphorbiaceae

(2)

UB1

VH

ypse

rpa

dec

um

ben

s(Benth)Diels

Menispermaceae

(4)

UB1G

MELU102316

VH

ypse

rpa

lau

rina(FM

uell)Diels

Menispermaceae

(2)

LM

Webb(1949)DNT

MELU102276

VH

ypse

rpa

smil

aci

foli

aDiels

Menispermaceae

(1)

UB2

TH

ypso

ph

ila

die

lsia

naLoes

Celastraceae

(3)

UB1

MELU102311

TIr

vin

gb

ail

eya

au

stra

lis(CTW

hite)

RAHoward

Icacinaceae

(2)

UB1G

MELU102324

TIx

ora

bifl

ora

Fosberg

Rubiaceae

(2)

LM

TIx

ora

sp(N

orthMaryLA

BP

Hyl

and

86

18)

Rubiaceae

(3)

UB1B2

VJa

smin

um

did

ymu

mGForst

Oleaceae

(2)

UB1B2G

VJa

smin

um

kaje

wsk

iiCTW

hite

Oleaceae

(2)

UB2G

SH

La

sia

nth

us

stri

gosu

sWight

Rubiaceae

(2)

LM

SHT

Lee

ain

dic

a(Burm

f)Merr

Leeaceae

(1)

LM

SHT

Lep

idoza

mia

ho

pei

(WHill)Regel

Zam

iaceae

(3)

LM

TL

eth

edo

nse

tosa

(CTW

hite)

Kosterm

Thymelaeaceae

(2)

UG

P(T)

Lic

ua

lara

msa

yi(FM

uell)Domin

Arecaceae

(3)

LM

P(SH)

Lin

osp

ad

ixm

inor(W

Hill)FM

uell

Arecaceae

(2)

LM

TL

itse

ab

ind

on

ianaFM

uell(FM

uell)

Lauraceae

(2)

UG

MELU102097

TL

itse

aco

nn

ors

iiBHyland

Lauraceae

(2)

UGR

TL

itse

ale

efea

na(FM

uell)Merr

Lauraceae

(7)

UL

B1B2M

MELU102145

TL

og

an

iace

aesp

Loganiaceae

(1)

UG

TL

om

ati

afr

axi

nif

oli

aFM

uellex

Benth

Proteaceae

(2)

UB2G

EE(Conn(perscomm)ndashve

L

sila

ifo

liaRBrftflwr+C

NWebb(1949)1948

MELU102091

TM

aca

ran

ga

inam

oen

aFM

uellex

Benth

Euphorbiaceae

(7)

UB1

Webb(1949)DNT

MELU102156

TM

aca

ran

ga

sub

den

tata

Benth

Euphorbiaceae

(1)

LM

SH

Ma

ckin

laya

con

fusa

Hem

sl

Araliaceae

(1)

UR

SH

Ma

ckin

laya

ma

cro

scia

dea

(FM

uell)FM

uell

Araliaceae

(4)

UB1G

Webb(1949)DNT

VM

aes

ad

epen

den

sFM

uell

Myrsinaceae

(1)

UR

SHT

Mel

ico

pe

bro

ad

ben

tia

naFM

Bailey

Rutaceae

(3)

UB1G

MELU102131

TM

elic

op

ejo

nes

iiTGHartley

Rutaceae

(1)

UB2

TM

elic

op

evi

tifl

ora

(FM

uell)TGHartley

Rubiaceae

(1)

UL

B1GM

VM

elo

din

us

acu

tifl

oru

sFM

uell

Apocynaceae

(1)

LM

Webb(1949)DNT

VM

elo

din

us

au

stra

lis(FM

uell)Pierre

Apocynaceae

(4)

UB1GR

VM

elodin

us

bacc

elli

anus(FM

uell)STBlake

Apocynaceae

(3)

UB1B2

MELU102230

VM

elo

do

rum

uh

riiFM

uell

Annonaceae

(1)dry

LM

TM

isch

ary

tera

laute

reri

an

a(FM

Bailey)HTurner

Sapindaceae

(2)

UB2GR

MELU102286

1040 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TM

isch

oca

rpu

sex

an

gu

latu

s(FM

uell)Radlk

Sapindaceae

++

UB1

BRI578801

MELU102190

102288

TM

isch

oca

rpu

sg

rand

issi

mus(FM

uell)Radlk

Sapindaceae

++

UL

GM

BRI578802

MELU102287

TM

isch

oca

rpu

sla

chn

oca

rpu

s(FM

uell)Radlk

Sapindaceae

(3)

UB2G

TM

isch

oca

rpu

sm

acr

oca

rpu

sSTReynolds

Sapindaceae

(3)

UB1B2

TM

isch

oca

rpus

pyr

iform

is(FM

uell)

Radlksubsp

p

yrif

orm

isSapindaceae

(1)

UB2

TMonim

iaceae

Gen(AQ63687)sp(D

avies

Creek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB2R

VM

ori

nd

aGen(AQ124851)sp

(Boonjie

LJ

Web

b+

68

37A)

Rubiaceae

(3)

UB1

VM

ori

nd

aja

smin

oid

esACunnex

Hook

Rubiaceae

(1)

UB2G

Webb(1949)ndashCN

VM

ori

nd

asp

Rubiaceae

(1)

UG

VM

ori

nd

au

mb

ella

taL

Rubiaceae

(1)

UG

TM

usg

rave

ah

eter

op

hyl

laLSSm

Proteaceae

(2)

LM

EEConn(perscomm)

1of8+v

eT

Musg

rave

ast

enost

ach

yaFM

uell

Proteaceae

(4)

UGR

EEConn(perscomm)ndashve

TM

yris

tica

glo

bo

sasubsp

mu

elle

ri(W

arb)WJdeWilde

Myristicaceae

(3)

UB1

TM

yris

tica

insi

pid

aRBr

Myristicaceae

(3)

LM

MELU102312

TN

eiso

sper

ma

po

wer

i(FM

Bailey)

Fosbergamp

Sachet

Apocynaceae

(2)

UB2

TN

eoli

tsea

dea

lba

ta(RBr)Merr

Lauraceae

(4)

UB1B2

Gibbs(1974)ndashCN

VN

eose

pic

aea

jucu

nda(FM

uell)Steenis

Bignoniaceae

(2)

LM

TN

iem

eyer

apru

nif

era(FM

uell)FM

uell

Sapotaceae

(4)

UL

B1GM

MELU102149

P(T)

No

rma

nbya

no

rman

byi

(WHill)LHBailey

Arecaceae

(4)

LM

TO

pis

thio

lep

ish

eter

op

hyl

laLSSm

Proteaceae

++

UB1B2

EE

Conn(perscomm)

flwrandlf+v

eBRI578808

MELU102298

P(T)

Ora

nio

psi

sa

pp

end

icula

ta(FM

Bailey)JDransf

AKIrvineamp

NW

Uhl

Arecaceae

(1)

UB1

VP

ach

ygo

ne

lon

gif

oli

aFM

Bailey

Menispermaceae

(2)

LM

TP

ala

quiu

mg

ala

cto

xylo

n(FM

uell)HJLam

Sapotaceae

(1)dry

LM

VP

alm

eria

sca

nden

sFM

uell

Monim

iaceae

(4)

UG

MELU102194

SHT

Pa

nd

an

us

mo

nti

cola

FM

uell

Pandanaceae

(1)

UG

VP

an

do

rea

pa

ndo

rana(A

ndrews)

Steenis

Bignoniaceae

(1)

UB2

Webb(1949)DNT

VP

ara

rist

olo

chia

au

stra

lop

ith

ecu

rus(FM

uell)

MichaelJParsons

Aristolochiaceae

(3)

UB1B2G

VP

ars

on

sia

lati

foli

a(Benth)STBlake

Apocynaceae

++

UB1GR

Webb(1949)DNT

other

Pa

rso

nsi

asppndashCN

BRI578800

MELU102128

VP

ars

on

siasp1

Apocynaceae

(1)

UG

VP

ars

on

sia

lan

gia

naFM

uell

Apocynaceae

(1)

UR

VP

ass

iflora

sp(K

uranda

BH

128

96)

Passifloraceae

++

LM

BRI578813

MELU102297

TP

erro

ttet

iaa

rbore

scen

s(FM

uell)Loes

Celastraceae

(1)

UG

SHT

Pil

idio

stig

ma

pa

pu

an

um

(Lauterb)AJScott

Myrtaceae

(3)

LM

SH

Pil

idio

stig

ma

tetr

am

erum

LSSm

Myrtaceae

(4)

UB1

MELU102274

TP

ilid

iost

igm

atr

op

icum

LSSm

Myrtaceae

(2)

UB1

VP

iper

can

inu

mBlume

Piperaceae

(3)

LM

MELU102265

VP

iper

no

vae-

ho

lla

nd

iaeMiq

Piperaceae

(3)

UB1G

Webb(1949)DNT

TP

ita

via

ster

ha

plo

ph

yllu

s(FM

uell)TGHartley

Rutaceae

(12)

UL

B1GM

MELU102100

102187

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1041

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022

APPENDIX

Co

nti

nu

ed

Lifeform

Species

Fam

ily

HCN

+ndashenz(n

tested)UplandlowlandSite(substrate)

Previousreports

Lodgem

entno

SHT

Pit

tosp

oru

mru

big

ino

sum

ACunn

Pittosporaceae

(3)

UL

B1B2GRM

P

un

dula

tum

(Bailey1909in

Webb1949)1948

MELU102192

TP

itto

spo

rum

win

giiFM

uell

Pittosporaceae

(2)

UG

TP

laco

sper

mum

cori

ace

um

CTW

hiteamp

WDFrancis

Proteaceae

(2)

UG

TP

od

oca

rpus

dis

per

mu

sCTW

hite

Podocarpaceae

(2)

UB1

TP

oly

alt

hia

mic

hael

iiCTW

hite

Annonaceae

DNT

UB1

TP

oly

osm

aa

lan

gia

ceaFM

uell

Grossulariaceae

(3)

UGR

TP

oly

osm

ah

irsu

taCTW

hite

Grossulariaceae

(1)

UB1B2

TP

oly

osm

arh

yto

ph

loia

CTW

hiteamp

WDFrancis

Grossulariaceae

(1)

UB2R

Webb(1949)DNT

TP

oly

osm

ari

gid

iusc

ula

FM

uellamp

FM

Baileyex

FM

Bailey

Grossulariaceae

(3)

UB1

TP

oly

scia

sa

ust

rali

ana(FM

uell)Philipson

Araliaceae

++

UL

B1B2GRM

BRI578812

MELU102301

TP

oly

scia

sm

oll

is(Benth)Harmsamp

CTW

hite

Araliaceae

(1)

UB1

TP

oly

scia

sm

urr

ayi

(FM

uell)Harms

Araliaceae

(1)

UB1

Webb(1949)DNT

SH

Po

lysc

ias

pu

rpu

reaCTW

hite

Araliaceae

(4)

UG

MELU102154

HE

Po

tho

slo

ng

ipes

Schott

Araceae

(4)

UL

B1M

TP

ou

teri

aa

ster

oca

rpo

n(PRoyen)Jessup

Sapotaceae

(1)

UB2

TP

ou

teri

ab

row

nle

ssia

na(FM

uell)Baehni

Sapotaceae

(3)

UL

B1B2GM

TP

ou

teri

aca

stan

osp

erm

a(CTW

hite)

Baehni

Sapotaceae

(4)

UB1G

TP

ou

teri

ach

art

ace

a(FM

uellex

Benth)Baehni

Sapotaceae

(3)2dry

LM

TP

ou

teri

am

yrsi

no

den

dro

n(FM

uell)Jessup

Sapotaceae

(7)

UL

GM

MELU102221

TP

ou

teri

ap

ap

yra

cea(PRoyen)Baehni

Sapotaceae

(2)

UB2

MELU102308

102309

TP

ou

teri

ap

ears

on

ioru

mJessup

Sapotaceae

(1)

UR

TP

runus

turn

eria

na(FM

Bailey)Kalkman

Rosaceae

++

UL

B1M

BRI578814

MELU102133

102134

HP

seu

der

an

them

um

vari

ab

ile(RBr)Radlk

Acanthaceae

(1)

LM

Webb(1949)DNT

TP

seu

du

vari

afr

og

ga

ttii(FM

uell)Jessup

Annonaceae

(3)

LM

MELU102310

TP

seu

du

vari

am

ulg

rave

anaJessup

var

gla

bre

scen

sAnnonaceae

(1)

UG

SH

Psy

chotr

iad

all

ach

ianaBenth

Sapindaceae

(1)

UB2

SH

Psy

chotr

ialo

nic

ero

ides

Sieber

exDC

Rubiaceae

(2)

UG

Webb(1949)DNT

SH

Psy

chotr

ian

ema

top

odaFM

uell

Rubiaceae

(2)

LM

SH

Psy

chotr

iasp

Rubiaceae

(1)

LM

SH

Psy

chotr

iasp(U

tchee

Creek

H

Fle

cker

NQ

NC

53

13)Rubiaceae

(3)

UG

SH

Psy

chotr

iasu

bm

on

tan

aDomin

Rubiaceae

(2)

UB1G

TP

syd

rax

laxi

flo

ren

sS

TR

eyn

old

samp

RJ

FH

end

Rubiaceae

(2)

UG

TP

ull

east

utz

eri(FM

uell)Gibbs

Cunoniaceae

(3)

UG

SH

Ra

ndia

sp(Boonjie

LW

Jes

sup+

GJM

26

4)

Rubiaceae

(1)

UB1

SH

Ra

ndia

tuber

culo

saFM

Bailey

Rubiaceae

(7)

UB1G

MELU102158

TR

ap

an

eaa

chra

dif

oli

a(FM

uell)Mez

Myrsinaceae

(3)

UB2GR

MELU102217

SHT

Ra

pan

eap

oro

sa(FM

uell)Mez

Myrsinaceae

(3)

UL

RM

SH

Ra

pan

easp(A

therton

KJ

Wh

ite

AQ

917

78)

Myrsinaceae

(2)

UR

MELU102260

HE

Rh

aph

ido

pho

rap

etri

eanaAHay

Araceae

(3)

LM

Webb(1949)DNT

TR

ho

da

mn

iab

lair

ian

aFM

uell

Myrtaceae

(2)

UG

TR

ho

da

mn

iasp

ong

iosa

(FM

Bailey)Domin

Myrtaceae

(2)

UR

1042 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

loco

sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

vio

ides

(FM

Bailey)BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

john

son

ii(FM

uell)BHyland

Myrtaceae

(3)

UB2G

TS

yzyg

ium

kura

nd

a(FM

Bailey)BHyland

Myrtaceae

(4)

UL

B1GM

TS

yzyg

ium

lueh

ma

nn

ii(FM

uell)LASJohnson

Myrtaceae

(1)

UR

TS

yzyg

ium

mo

no

sper

mu

mCraven

Myrtaceae

(1)

LM

TS

yzyg

ium

pa

pyr

ace

um

BHyland

Myrtaceae

(3)

UB1G

MELU102159

TS

yzyg

ium

trach

yph

loiu

m(CTW

hite)

BHyland

Myrtaceae

(1)

UB1

TS

yzyg

ium

wes

aBHyland

Myrtaceae

(3)

UB2G

MELU102096

102249

TS

yzyg

ium

wil

son

iisubsp

cryp

top

hle

biu

m(FM

uell)BHyland

Myrtaceae

(3)

UB1G

MELU102223

SH

Taber

naem

onta

na

pandaca

quiLam

Apocynaceae

(1)

LM

SH

Ta

pei

no

sper

masp(Cedar

Bay

JG

Tra

cey

14

78

0)

Myrsinaceae

(4)

UG

SH

Ta

sma

nn

iain

sip

idaRBrex

DC

Winteraceae

(3)dry

UR

TT

ernst

roem

iach

erry

i(FM

Bailey)

Merrex

JFBaileyamp

CTW

hite

Theaceae

(2)

LM

VT

etra

cera

no

rdti

an

avar

no

rdti

an

aFM

uell

Dilleniaceae

(2)

UL

GM

TT

etra

syn

an

dra

laxi

flo

ra(Benth)JRPerkins

Monim

iaceae

(6)

UL

B1GM

MELU102229

TT

imo

niu

ssi

ng

ula

ris(FM

uell)LSSm

Rubiaceae

(1)

UB1

TT

oec

him

aer

yth

roca

rpu

m(FM

uell)Radlk

Sapindaceae

(3)

UL

B1GM

MELU102321

TT

oec

him

am

on

tico

laSTReynolds

Sapindaceae

(1)

UGR

MELU102098

TT

riu

nia

eryt

hro

carp

aForeman

Proteaceae

(3)

UB1

EEConn(perscomm)ndashve

VT

rop

his

sca

nden

s(Lour)Hookamp

Arnsubsp

sc

an

den

sMoraceae

(1)

LM

MELU102271

VU

nca

ria

lan

osa

var

ap

pen

dic

ula

ta(Benth)Ridsdale

Rubiaceae

(1)

LM

TU

rom

yrtu

sm

etro

sider

os(FM

Bailey)AJScott

Myrtaceae

(1)

UR

MELU102315

VV

enti

lag

oec

oro

llata

FM

uell

Rham

naceae

DNT

LM

TW

ilki

eaa

ng

ust

ifoli

a(FM

Bailey)JRPerkins

Monim

iaceae

(1)

UB1

MELU102330

SH

Wil

kiea

sp

Monim

iaceae

(2)

UB1

SH

Wil

kiea

sp(Barong

LW

Jes

sup

71

9)

Monim

iaceae

(14)

UB1B2G

SH

Wil

kiea

spGen(A

Q63687)sp

(DaviesCreek

LJ

Web

b+

64

30)

Monim

iaceae

(3)

UB1

MELU102302

SH

Wil

kiea

ward

elli

i(FM

uell)JRPerkins

Monim

iaceae

(3)

UR

TX

an

thop

hyl

lum

oct

an

dru

m(FM

uell)Domin

Xanthophyllaceae

(3)

UL

B1B2GM

MELU102101

TX

ylop

iam

acc

rea

e(FM

uell)LSSm

Annonaceae

(1)

LM

TZ

anth

oxy

lum

ven

eficu

mFM

Bailey

Rutaceae

(2)

UB2G

Webb(1949)DNT

MELU102215

daggerSpeciesin

thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TR

hodom

yrtu

sper

vagata

Guymer

Myrtaceae

(3)

UB2GR

TR

hys

oto

echia

mo

rto

nia

na(FM

uell)Radlk

Sapindaceae

(1)

UB2

TR

hys

oto

echia

rob

erts

onii(FM

uell)Radlk

Sapindaceae

(1)

LM

VR

ipo

go

nu

ma

lbu

mRBr

Smilacaceae

(3)

UB1B2G

VR

ipo

go

nu

mel

seya

nu

mFM

uell

Smilacaceae

(3)

UB1

VR

ou

rea

bra

chya

nd

raFM

uell

Connaraceae

(1)

LM

SH

Rubus

quee

nsl

andic

usARBean

Rosaceae

(1)

UB2

TR

ypa

rosa

java

nica(Blume)

Kurz

exKoordamp

ValetonD

Achariaceae

dagger++

LM

Pam

mel

(1911)

R

caes

ia+C

N

Rosenthaler

(1919)

Ryp

aro

saspp+C

N

MELU102277

VS

arc

op

eta

lum

ha

rvey

anu

mFM

uell

Menispermaceae

(1)

BB1

Webb(1949)DNT

Hurst(1942)ndashCN

TS

arc

op

tery

xm

art

yan

a(FM

uell)Radlk

Sapindaceae

DNT

UG

TS

arc

op

tery

xre

ticu

lata

STReynolds

Sapindaceae

(2)

LM

TS

arc

oto

echia

lan

ceo

lata

(CTW

hite)

STReynolds

Sapindaceae

(2)

UG

TS

arc

oto

echia

pro

tra

ctaRadlk

Sapindaceae

(3)

UB1G

TS

arc

oto

echia

sp(M

tCarbine

LW

Jes

sup+

GJM

99

5)

Sapindaceae

(2)

UR

MELU102303

VS

caev

ola

ena

nto

ph

ylla

FM

uell

Goodeniaceae

(4)

UB1G

Webb(1949)DNT

TS

chis

toca

rpa

eajo

hn

soniiFM

uell

Rham

naceae

(5)

UB1

TSch

izom

eria

whit

eiMattf

Cunoniaceae

(1)

UG

TS

colo

pia

bra

un

ii(K

lotzsch)Sleumer

Salicaceaedagger

(1)

UB2

TS

iph

on

od

on

mem

bra

na

ceu

sFM

Bailey

Celastraceae

(3)

UL

B1M

TS

loa

nea

au

stra

lissubsp

pa

rvifl

ora

Coode

Elaeocarpaceae

(4)

UB1

TS

loa

nea

lan

giiFM

uell

Elaeocarpaceae

(5)

UL

B2GM

TSlo

anea

macb

rydei

FM

uell

Elaeocarpaceae

(3)

UB1G

VS

mil

ax

acu

lea

tiss

imaConran

Smilacaceae

(3)

UB1

VS

mil

ax

calo

ph

ylla

Wallex

ADC

Smilacaceae

(1)

LM

VS

mil

ax

gla

uca

Walter

Smilacaceae

(2)

UB1

VS

mil

ax

gly

cip

hyl

laSm

Smilacaceae

(2)

UB2R

SH

So

lan

um

ma

coo

raiFM

Bailey

Solanaceae

(1)

UB2

SH

So

lan

um

sp

Solanaceae

(2)

UB1

SH

So

lan

um

viri

dif

oli

um

Dunal

Solanaceae

(1)

UG

TS

ph

eno

stem

on

lob

osp

oru

s(FM

uell)LSSm

Sphenostem

onaceae

(3)

UGR

MELU102111

SHT

Ste

ga

nth

era

au

stra

lia

naCTW

hite

Monim

iaceae

(2)

UB2

TS

tega

nth

era

ma

coo

raia

(FM

Bailey)PKEndress

Monim

iaceae

(4)

UR

MELU102231

TSte

noca

rpus

reti

cula

tusCTW

hite

Proteaceae

(1)

UR

EEConn(perscomm)ndashve

MELU102196

TS

teno

carp

us

sin

ua

tus(Loudon)Endl

Proteaceae

(1)flwrndash

UB2

EEConn(perscomm)ndashve

VS

teph

an

iaja

po

nic

a(Thunb)Miers

Menispermaceae

DNT

LM

TSto

rcki

ella

aust

rali

ensi

sJHRoss

ampBHyland

Caesalpiniaceae

(3)

LM

MELU102279

TS

treb

lus

gla

ber

var

a

ust

rali

anu

s(CTW

hite)

Corner

Moraceae

(3)

UB2R

VS

tryc

hn

os

min

orDennst

Strychnaceae

(1)

LM

TS

ymp

loco

sco

chin

chin

ensi

ssubsp1

Symplocaceae

(2)

UG

TS

ymp

loco

sco

chin

chin

ensi

svar

g

itto

nsi

iNoot

Symplocaceae

(2)

UB2R

TS

ymp

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sco

chin

chin

ensi

svar

pil

osi

usc

ula

Noot

Symplocaceae

(3)

UB2GR

MELU102218

SH

Sym

plo

cos

ha

yesi

iCTW

hiteamp

WDFrancis

Symplocaceae

(3)

UB1B2R

MELU102320

SHT

Sym

plo

cos

pa

uci

sta

min

eaFM

uellamp

FM

Bailey

Symplocaceae

(3)

UL

B1GM

TS

ynim

aco

rdie

roru

m(FM

uell)Radlk

Sapindaceae

(1)

UL

B2GM

TS

ynim

am

acr

op

hyl

laSTReynolds

Sapindaceae

(4)

UB1B2G

MELU102289

TS

yno

um

mu

elle

riCDC

Meliaceae

(2)

UB2

MELU102306

TS

yzyg

ium

can

ico

rtex

BHyland

Myrtaceae

(2)

UB2G

MELU102272

Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests 1043

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icoupcomaobarticle9761017206982 by guest on 11 January 2022

TS

yzyg

ium

corm

iflo

rum

(FM

uell)BHyland

Myrtaceae

(3)

UL

B1B2GM

MELU102224

TS

yzyg

ium

end

oph

loiu

mBHyland

Myrtaceae

(4)

UB1B2GR

MELU102153

TS

yzyg

ium

gu

sta

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UR

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pei

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cey

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rea

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aro

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ed(W

ebber2005)

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limited

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TS

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ium

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iaceae

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oec

him

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roca

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him

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Wil

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sup

71

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Wil

kiea

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kiea

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oxy

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ven

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thefamiliesSalicaceaeandAchariaceae

whichuntilrecentrevisionswerein

theFlacourtiaceae

(Chase

eta

l2002)

zNon-w

oody

Alo

casi

ab

risb

anen

siswas

notincluded

intheanalysisofthefrequency

ofcyanogenic

species

DR

ypa

rosa

java

nic

aiscurrentlythesubject

oftaxonomic

reviewthisQueensland

Ryp

aro

saspislikelyto

berenam

ed(W

ebber2005)

Dueto

limited

access

tofoliagetestsforcyanogenesisusingfreshtissuewerenotconductedinsteadfreeze-dried

sampleswereassayed

quantitativelyforthepresence

ofCNindicated

aslsquodryrsquo

1044 Miller et al mdash Frequency of Cyanogenesis in Australian Tropical Rainforests

Dow

nloaded from httpsacadem

icoupcomaobarticle9761017206982 by guest on 11 January 2022