Habitat-use by the Green and Golden Bell Frog Litoria aurea in Australia and New Zealand

12 April 2002Australian

Zoologist volume 32 (1)

Habitat-use by the Green and GoldenBell Frog Litoria aurea in Australia andNew ZealandGraham H.Pyke1 Arthur W.White1,Phillip J.Bishop2,Bruce Waldman3

1.Frog Ecology and Behaviour Group Australian Museum 6 College St. Sydney NSW 2000 Australia2.Department of Zoology University of Otago PO Box 56 Dunedin New Zealand3.Department of Zoology University of Canterbury Private Bag 4800 Christchurch New ZealandEmail for corresponding author: [email protected]

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Patterns of habitat-use are used widely as management guides in theconservation of wildlife. However, even for relatively well-studied species, suchthe Green and Golden Bell Frog Litoria aurea, our knowledge of specific habitatrequirements is lacking.This study sought to compare the patterns of breeding habitat shown by L. aureain New South Wales (NSW), Victoria and New Zealand (where it is a feralspecies), with those described in an earlier study of Pyke and White (1996), toreview the conservation status of this species in Victoria,and to relate its observedpatterns of distribution and habitat-use to the distribution of the introducedPlague Minnow (Gambusia holbrooki in Australia; G. affinis in New Zealand).We found that L. aurea used similar breeding habitats in NSW,Victoria and NewZealand.Across these areas its breeding is almost completely restricted to waterbodies that are still, relatively unshaded, and low in salinity (i.e., <7.3 ppt).All ofits known breeding sites are highly disturbed,mostly through human activities butalso through flooding and other natural processes. It generally breeds in small(i.e., <1000 sq m), shallow (i.e., <1 m) ponds that are either ephemeral orfluctuate significantly in water level,are free of Gambusia and other predatory fish,and have emergent aquatic vegetation. Its breeding habitat also usually haspotential shelter provided by nearby rocks or thick, low vegetation.Yet L. aurea is adaptable to different habitats and we found it breeding in a widerange of conditions including ponds lacking emergent vegetation and thosealready colonised by Gambusia. Breeding ponds ranged in terms of substrate,nearby terrestrial vegetation, water source, and water properties includingturbidity, dissolved oxygen, oxidation reduction potential, pH and temperature.Gambusia may have had a negative impact on the distribution of L. aurea in NewZealand and could threaten populations in Victoria.We failed to find L.aurea atsites in New Zealand where it had previously been reported but that are nowcolonised by Gambusia.This fish has not invaded L. aurea habitat in Victoria, butit can potentially do so from nearby regions. Further surveys for L. aurea inVictoria and New Zealand seem warranted.Paradoxically New Zealand may offer unique opportunities for furtherresearch on the biology of Australian frogs.The commonly encountered frogsin New Zealand are three Australian species, two of which (i.e., L. aurea and L.raniformis) are considered threatened with extinction in Australia. It may bepossible in New Zealand to study relatively large populations of theseAustralian frog species, in relatively simple frog communities and habitats.Because these species are not protected in New Zealand, field studies can alsoincorporate experimental manipulations not readily possible in Australia.

Key words: Australia, New Zealand, frog, Litoria aurea, Litoria raniformis, habitat, Gambusia, Plague Minnow,Mosquito Fish

13April 2002Australian


IntroductionKnowledge of patterns of habitat-use is necessaryto understand, conserve and manage animalpopulations. Understanding how animals chooseand make use of habitat allows us to formulateplans to facilitate their survival and reproduction.Moreover, this information is necessary for thedevelopment of accurate survey strategies (e.g.,Soulé 1986). The compilation of habitat-useinformation thus represents an important firststep in the conservation and management ofspecies that are considered to be undersignificant risk of extinction (i.e., classified as‘threatened’, ‘endangered’, or ‘vulnerable’).

Despite their importance, however, patterns ofhabitat-use have been little documented for frogsin general, and for frogs in Australia and NewZealand in particular. Most descriptions ofhabitat for Australian frogs focus on the rates ofwater flow and the nature of adjacent or nearbyvegetation (e.g., Barker et al. 1995; Robinson1993; Cogger 2000). Habitat descriptions forNew Zealand frogs generally focus on broadvegetation categories, distance from water,presence/absence of shade, presence/absence ofrocks and elevation range (e.g., Bell 1982a; Gilland Whitaker 1996; Newman 1996).

The most detailed studies of patterns of habitat-use for frogs of Australia and New Zealand havebeen carried out for three Australian species,Litoria aurea, L. spenceri and L. pearsoniana(Gillespie and Hollis 1996; Pyke and White 1996;Parris 2001). In an earlier study we used stepwiselogistic regression to compare sites where breedingby L. aurea had been recorded (i.e., tadpoles andmetamorphs observed) with sites where thisspecies was known to be present, but where breed-ing had not yet been recorded (Pyke and White1996). We found that L. aurea usually breeds inwater bodies that are still, shallow, unshaded, freeof predatory fish such as the introduced PlagueMinnow Gambusia holbrooki (also known asGambusia or Mosquito Fish), and ephemeral(rather than having fluctuating or constant waterlevel), with sand or rock substrate and withemergent aquatic plants (Pyke and White 1996).We also found that shelter is usually providedthrough nearby rock or thick, low vegetation, thatadjacent and nearby vegetation may vary fromgrassland to heath/shrubland to woodland but hasnot so far included forest, and that many of thesites where L. aurea breeds are highly disturbed byhuman activities (Pyke and White 1996).

Gillespie and Hollis (1996) used contingencytable and Spearman rank correlation analysis todetermine correlations between variousquantified measures of stream habitat and bothpresence/absence of L. spenceri and its relativeabundance (i.e., number adults observed per kmlength of stream). They found that L. spencerioccurred almost exclusively in association withrock habitats along streams, and that itsabundance was negatively correlated withclearance of adjacent forest, human access andthe presence of trout (Gillespie and Hollis 1996).Parris (2001) used stepwise logistic and linearregression to determine relationships betweenvarious climatic and habitat variables and twomeasures of abundance for L. pearsoniana (i.e.,presence/absence and log transformed number ofdetected individuals for those sites where specieswas present). She found that L. pearsoniana wasmost likely to occur at large streams with mesicmid-storey vegetation in riparian zone, asindicated by presence of palms, and that, at siteswhere this species was present, its abundanceincreased with increasing stream size (Parris2001). Similar, though less detailed, studies ofhabitat-use for Australian frogs have been carriedout by Healey et al. (1997) and Lemckert (1999).

Even for L. aurea, L. spenceri and L. pearsoniana,however, our understanding of habitat-use remainslimited. Studies of L. aurea habitat have not, forexample, assessed water chemistry (e.g., pH,salinity, contaminants), nor physical characterisics(e.g., temperature, turbidity). How habitat-usemight vary with time or location has not beenconsidered (Pyke and White 1996). Pyke andWhite (1996) considered sites only within NSWand did not consider other areas where L. aureaalso occurs. Studies of habitat-use by L. spenceri andL. pearsoniana similarly omitted physical andchemical properties of available water and wereunable to provide specific detailed information inrelation to breeding habitat (Gillespie 1992;Gillespie and Hollis 1996; Parris 2001). The rangesof habitats at sites where these two species occurare likely to be considerably broader than thesubsets of breeding sites, as is the case for L. aurea(e.g., Gillespie 1996; Pyke and White 1996).

Litoria aurea is endemic to Australia, but it hasbeen introduced to a number of islands in theSouth Pacific (see Pyke and White 2001). InNew Zealand, for example, it occurs in manylocations in the northern part of the North Island(Thomson 1922; McCann 1961; Bull andWhitaker 1975; Bell 1982a,b; Pickard and Towns



Pyke et al.

1988). It has also been introduced into NewCaledonia, Vanuatu, the Loyalty Islands and theislands of Wallis and Futuna, and may be locallyabundant in these feral populations (e.g.,McCann 1961; Gibbons 1985; Tyler 1976, 1979;Bell 1982a; Bauer et al. 1997; Bauer and Sadlier2000). No comparisons have been made betweenits patterns of habitat-use where introduced withthose it shows within its endemic range

Within Australia, the legal conservation status ofL. aurea and the factors apparently affecting itspopulations vary between NSW and Victoria. InNSW, it is considered ‘endangered’ (i.e., underthe Threatened Species Conservation Act 1995),having disappeared from over 80% of its originaldistribution and with many of its remainingpopulations being small (White and Pyke 1996;White 1997). However, in Victoria, declines inthe species have not been so apparent, and it isnot presently considered as either ‘vulnerable’ or‘endangered’ (i.e., under the Flora and FaunaGuarantee Act 1988) (Gillespie 1996).

The dramatic decline in the distribution andabundance of L. aurea in NSW appears to haveresulted from habitat modification anddestruction, predation on eggs and tadpoles bythe introduced G. holbrooki, and possibly disease(e.g., Mahony 1993, 1999, 2001; Daly 1995;Morgan 1995; Goldingay 1996; Mahony 1996;Morgan and Buttemer 1996; Osborne et al. 1996;Pyke and White 1996; Gillespie 1997; Tyler1997; Goldingay and Lewis 1999). In the north-east Gippsland area of Victoria, on the otherhand, most of the sites where L. aurea occurs arefree of Gambusia. This fish is abundant to boththe north and west of this region (Cadwalladerand Backhouse 1983; Gillespie 1996; Pyke andWhite, unpublished). Perhaps L. aurea hasdeclined less in Victoria because of the absence ofG. holbrooki. No comparison has so far been madebetween patterns of its habitat-use in areas whereit has suffered pronounced decline and those inareas where such declines are not so evident.

The aims of the present study were therefore tocompare the patterns of habitat-use shown by L.aurea in NSW, Victoria and New Zealand; tocompare the patterns of habitat-use determinedin this study with those described in the earlierstudy of Pyke and White (1996); to review theconservation status of this species in Victoria;and to relate its observed patterns ofdistribution and habitat-use to the distributionof the Plague Minnow.

MethodField work for this study was carried out in NSW,Victoria and New Zealand. We used similarmethods in Victoria and New Zealand, and sodescribe those together, with methods for NSWdescribed separately.

For all three regions, recorded frog locationswithin about 500 m of one another wereconsidered to represent a single “site” asmovements of up to 200-300 m are oftenrecorded for this species, whereas movements ofgreater than 500 m are rarely detected (Pyke andWhite 2001). The minimum recorded distancebetween such “sites” was at least 5 km.

The Plague Minnow is one species, G. holbrooki,in Australia and another closely related species,G. affinis, in New Zealand (McDowall et al. 1974;McDowall 1978; Lloyd and Tomasov 1985). Weassume that these two species are ecologicallyequivalent and use the genus name Gambusia torefer collectively to them.

All four authors participated in the New Zealandcomponent of this study, whereas the Australiancomponents involved just GP and AW. In bothcountries a number of volunteers assisted.A. Victoria and New ZealandThe Victorian and New Zealand componentsinvolved field trips, one to north-east Gippslandin Victoria, Australia (11-18 December 1996)and the other to the North Island of NewZealand (22-29 January 1997). In both casesvisits were made to as many as possible of thesites where L. aurea had previously been found.We surveyed these sites for the presence of frogsand at each we recorded a number of habitatvariables for each site. No rain fell during eitherof these field trips and temperatures weregenerally mild to warm (i.e., 15-25 0 C).

We visited 41 of 97 identified L. aurea sites inVictoria and 12 of 31 in New Zealand (Tables 1-2).We first identified sites where the species had beenrecorded using information that was supplied byvarious Government agencies and from personalcontacts or responses to media requests(Appendix). Logistical considerations thendetermined the sites that we were able to visit(Tables 1-2). Nevertheless, sufficient sites werevisited to be able to assess patterns of habitat use.

We adjusted the time spent at each site to ensurea reasonable likelihood of detecting L. aurea if itwere present, while allowing us to visit severalsites each day. We thus spent only about 30



minutes at sites where there was no open waterpresent, but from one to four hours at sites withwater, depending on the extent of potentialhabitat. Almost all surveys were carried outduring daylight hours, although we visited onesite in New Zealand also during the night. Giventhis regime, there was considerable variation inthe times of day when sites were visited. At eachsite there were at least five people involved in thesurvey. With this survey procedure we detectedL. aurea at 26% of sites in Victoria and NewZealand that had water present (see below).

At each site we visually searched for frogs sitting onreeds or other vegetation, floating in water orpresent under shelter. We also listened forresponses to call imitations. Any frogs seen orheard were identified to species. Hand nets (about14 x 20 cm in area, with a 40 cm long handle and2 mm mesh) were used to survey for tadpoles andGambusia whenever water was present. These weredragged for about 1 m through the water at variousdepths, in open water, around emergent vegetation

and under overhanging vegetation with eachsituation being thus surveyed at least twice. Anycaptured tadpoles were identified to species andthen released. Any captured Gambusia were notedand then released. In addition, we visuallyinspected areas of shallow water adjacent to thebank for the presence of Gambusia.

We visited all sites during spring and summerwhen reproductively active adults of L. aureacongregate around breeding ponds (Dankers1977; Brook 1980; Bell 1982a; Murphy 1994).During this period both males and females of thisspecies are often observed during the day, eitherin water or sitting on emergent vegetation. Inaddition, males of this species often call duringthe day, especially in response to either playing orimitating their call. We should have detectedtadpoles of this species if present, because theyusually occur in shallow water swimmingconspicuously just below the surface and areeasily captured with nets (Anstis 1974; Daly1995). We should similarly have detected

Figure 1.Locations of sites in Australia and New Zealand where L. aurea was recorded during this study. Siteswhere frog was found in essentially natural habitat are labelled.

Habitat-use by the Green and Golden Bell Frog



Pyke et al.

Gambusia whenever present, because they toousually occur in shallow water near the surfaceand typically are very abundant (Pollard et al.1980; Merrick & Gunther 1984; Allen 1989;Webb & Joss 1997). The time of day duringdaylight hours is not known to influence theresults of such surveys.

For each water body we recorded the same generalhabitat variables as Pyke and White (1996), butwith two modifications. We previously used theterm “permanent” to refer to water bodies withrelatively constant water level (Pyke and White1996), but this is misleading as many peopleconsider water bodies that fluctuate in water levelsbut never dry out as “permanent”. In the presentstudy, we recorded water level as being constant(C), fluctuating (F) or ephemeral (E). In addition,we previously recorded only the presence of visiblesigns of pollution, but this approach ignores thelikelihood of water pollution resulting from land-use practices within the water catchment. Hencewe also recorded any adjacent or nearby land-usepractices (such as grazing of livestock) that wouldbe expected to be sources of water pollution andwere upstream of the water body. Where suchsituations occurred we considered that pollution ofthe water was likely.

We also recorded a number of physical andchemical properties of each water body using aYeo-kal Series 6000 Water Meter. These includedturbidity (nts units), dissolved oxygen (g/litre),oxidation reduction potential (millivolts), salinity(parts per thousand - ppt), pH, conductivity andwater temperature.

At sites where we found no water body we recordedany areas within 50 m that showed signs ofaccumulating water after rain. We inferred thatdepressions with bottom mud or moist soil, withreeds or other emergent aquatic plants, or withdeposited material indicating a previous water lineconstituted potential breeding sites, as L. aureaoviposits in semi-permanent or ephemeral waterbodies (e.g., Pyke and White 1996, 2001).B. NSWThe third component of the study involved visits to11 known L. aurea sites in NSW that were notincluded in the earlier study of Pyke and White(1996) (Table 3). All but one of these sites wereunknown to us prior to 1994/95 when the earlierstudy was carried out. The Broughton Island site,though recorded earlier, was omitted for logisticalreasons. The visits to these 11 sites either spanneda full year or more at some sites, or occurred duringthe spring/summer breeding season of L. aurea (i.e.,September to February) (Table 3). They were all

carried out between February 1997 and February2001 (Table 3). During each visit we recorded thesame information as above.

The final component involved recording just thephysical and chemical properties of the water andthe presence/absence of L. aurea tadpoles at one ofthe NSW sites (Kurnell South) that was includedin Pyke and White (1996) (Table 3). This site wasincluded in the present study because the earlierstudy did not include these physical and chemicalwater properties, and because frequent visits havenow been made to this site at all times of year overa 2-3 year period, enabling water properties to bemeasured on many occasions when L. aureatadpoles have been present (Table 3).C. AnalysisA site was considered to be a breeding site for L.aurea if we either heard one or more males of thisspecies calling, observed tadpoles of the species, orobserved five or more adults including individualsof both sexes, with males having dark nuptial pads(an indication of current reproductive activity;Hamer 1998; Pyke and White 2001). It is unlikelythat frogs could have bred elsewhere and thenmoved into the site because there was never anyother potential breeding area within about 500m,L. aurea rarely moves more than about 500m, andthey are not known to move in groups (Pyke andWhite 2001).

We compared patterns of habitat-use that wereobserved for breeding in Victoria, New Zealandand NSW, and then compared patterns recorded inthis study with those previously observed for otherNSW sites (Pyke and White 1996). We did notcompare breeding and non-breeding sites, as inPyke and White (1996), because all but one of thesites where we detected L. aurea were considered tobe breeding sites (see below).We compared theproportions of breeding sites with particular habitatfeatures. As assumptions of the chi-squaregoodness of fit test were violated, we computedexact probabilities using 5000 iterations of theMonte Carlo simulation technique proposed byRoff and Bentzen (1989). We also compared theobserved ranges for each habitat parameter amongthe three areas. For the purposes of comparison wehave included tabulations of data collected as partof the earlier study by Pyke and White (1996).

ResultsOur degree of success in detecting L. aurea variedamong locations in Victoria, New Zealand andNSW. Litoria aurea was found at 4 of 41 sitesvisited in Victoria, 6 of 12 in New Zealand, andall of the 11 new sites in NSW.



In New Zealand our likelihood of detecting L.aurea at the sites we visited was lower whenGambusia were present. Litoria aurea weredetected in 6 of 10 water bodies that apparentlylacked Gambusia, but were not detected in any ofthree water bodies where we found Gambusia (P= 0.07). Litoria aurea had been recorded fromthese latter water bodies between about 6 and 12years earlier (Table 2). We did not find Gambusiaat any sites in Victoria, so L. aurea presence orabsence in habitats there must be attributed toother factors (Table 1).

Other habitat features also influenced ourlikelihood of detecting L. aurea. Litoria aurea weredetected only at sites with still, open water, andnot at any of the 15 Victorian sites that lackedwater at the time of our visit, nor at any of thefive Victorian sites that had only flowing water(Table 1). Still, open water was present at all ofthe sites in New Zealand and NSW that werevisited during this study (Tables 3-4).

In New Zealand, L. aurea was often the only frogspecies present at a site (Table 2), whereas inAustralia L. aurea was generally part of acommunity of several frog species (Tables 1 and 3;Pyke and White 1996). Of the five sites in NewZealand where we found L. aurea, L. raniformiswas also found at one site and L. ewingii was foundat another site (Table 2). Other frog species thatwe found along with L. aurea at the four breedingsites in Victoria included Crinia signifera (2 sites),Limnodynastes dumerilii (1 site) and Litoria lesueuri(1 site) (Table 1). In NSW L. aurea breeding sitesare generally shared with Crinia signifera and/orLimnodynastes peronii (Table 3; Pyke and White1996). As many as five other frog species may co-occur with L. aurea at its breeding sites inAustralia (Table 3; Pyke and White 1996).

At all but one of the 21 sites where we detected L.aurea, we also found evidence of breeding. All ofthe sites in Victoria and NSW, and all but one ofthe New Zealand sites, were considered to bebreeding sites (Tables 1-3).

Across these 20 breeding sites there was littlevariation in some of the general habitat parameters.All had still water that was either unshaded or onlypartially shaded (Table 5). Only one had Gambusiapresent, all but one were either ephemeral orfluctuating in water level, all but one had a sand orclay substrate, and all but one included thick, lowvegetation as potential shelter (Table 5). Sixteenwere largely or completely human-created, all butone of which was judged to receive high levels ofhuman disturbance, and sixteen had emergentaquatic vegetation (Table 5).

In contrast, the 20 sites varied substantially in theirother general habitat variables. Many water bodieswere, for example, less than 50 cm in maximumdepth, but a similar number were greater than 1 mdeep (Table 5). Although most water bodies wereless than 1000 sq m, some were larger (Table 5). Atmany sites nearby terrestrial vegetation includedbare ground, grassland or forest, and at some sitesshrubland and/or woodland of intermediate heightwas present nearby (Table 5). Sites where waterpollution was considered unlikely and sites where itwas considered likely were about equallyrepresented (Table 5).

Variation across the 20 breeding sites was veryhigh in all but one of the measured physical andchemical water properties. Observed ranges were0 to 148 nts for turbidity, 3 to 16 g/l for dissolvedoxygen, -531 to +351 millivolts for oxidationreduction potential, 3.7 to 9.8 for pH and 12 to37 deg. C for water temperature (Table 6). Onlyfor salinity was there relatively little variationwith an observed range of 0 to 7.3 ppt (Table 6),which are all low compared with sea-water whichhas a salinity of about 34 ppt (Pyke unpublished).

There were few apparent differences between theL. aurea breeding habitats across the three areas.In NSW, ephemeral and fluctuating water bodieswere about equally prevalent, whereas almost allthe breeding sites in Victoria and New Zealandhad fluctuating water bodies (Table 5). In NewZealand water sources were located in grazingland, natural vegetation or both, whereas inVictoria and NSW there were also other kinds ofwater sources (Table 5). Most of the NewZealand sites were considered likely to receivepolluted water, whereas most of the Victorian andNSW sites were considered to be unpolluted(Table 5). Potential shelter consisted only ofthick, low vegetation at the Victorian and NewZealand sites, whereas other kinds of shelter weresometimes available in NSW (Table 5).

Furthermore, the observed habitats at thebreeding sites in the present study were similar inmost respects to those that were considered inthe earlier study of Pyke and White (1996). Theproportions of the various habitat conditionswere similar between the two studies in terms ofthe degree to which a site was natural vs artificial,substrate, shade, water flow, water pollution,emergent aquatic plants, potential shelter, thenature of nearby terrestrial vegetation, andpresence/absence of Gambusia (Table 5), andnone of the observed differences between the twostudies for these habitat parameters wassignificant (P’s>0.05).




Pyke et al.

In addition, disturbance was found to be auniversal feature of the breeding habitat of L.aurea. Across all three regions and both studies, 35of 39 recorded L. aurea breeding sites wereartificial habitats that were subject to a high levelof anthropogenic disturbance (Table 5; Gillespie1996). We found evidence of L. aurea breeding atonly four essentially natural sites in Victoria andNSW (i.e., Victoria: Thurra River; NSW: LakeMeroo, Lake Conjola (Buckley’s Point) andBroughton Island; Tables 1, 3 and 5), none in NewZealand (Tables 2 and 5), and none in our earlierstudy (Pyke and White 1996). The four naturalsites also experience disturbance through flooding,drying-out or inundation with sea-water. The sitesat Lake Conjola (Buckleys Point), ShoalhavenHeads, Thurra River and Lake Meroo are knownto be affected by flooding (White unpublished),while the ponds on Broughton Island occasionallyeither dry out or are inundated by sea-water duringstorms (Pyke unpublished). Hence, all of theknown breeding sites of L. aurea experience highlevels of disturbance, either naturally or throughhuman activities.

A few significant differences emerged betweenthe two studies. In the earlier study, there weremore breeding sites with ephemeral water bodiesthan with fluctuating water levels, whereas thereverse held for the present study (Table 1; P =0.002). In the present study most sites receivedwater from natural vegetation, whereas few of thesites in the earlier study did so (Table 1; P =0.01). Also water bodies at breeding sites tendedto be shallower in the earlier study than in thepresent study (Table 1; P = 0.02).

DiscussionThe differences between NSW, Victoria and NewZealand in the total numbers of recorded sites forL. aurea, and how successful we were at detectingthis species when we revisited sites, may reflectdifferences in the nature and extent of previoussurvey effort rather than differences indistribution and/or abundance of the species. ForNSW, the total number of recorded locations forL. aurea (including many where it no longeroccurs) is now at least 191 with 163 having beenidentified in White and Pyke (1996), a further 11identified in the present study, plus at leastanother 7 having been discovered during the lasttwo to three years (R. Wellington, NSWNational Parks and Wildlife Service, pers. comm.2001). Here there has been considerable surveyeffort targeting L. aurea over an extended periodof time (e.g., Pyke and Osborne 1996). The 11NSW sites added during this study were the

result of recent, targeted surveys that indicatedthe presence of breeding populations of thespecies (e.g., Bannerman 1998; Pyke 1999). Onthe other hand, many of the 97 Victorian L. aureasites have resulted from recent opportunisticobservations rather than targeted survey andabout 88% were based on single frogs (Table 1;Gillespie 1996; pers. comm.). Furthermore, manywere based on observations of frogs found onroads during wet weather and hence may havebeen of dispersing frogs that were not necessarilyclose to breeding sites (G. Gillespie, VictorianDepartment of Natural Resources andEnvironment, pers. comm. 1997). Such frogswere unlikely to be detected when we revisitedthe sites during dry weather.

We have identified only 31 L. aurea sites in NewZealand, but there has been relatively little interestin the species here and no targeted surveys(Bishop and Waldman unpublished). Therecognised geographic range of the species in NewZealand is relatively small, being restricted to thearea north of about Rotorua in the North Island,although occasional unconfirmed reports havebeen obtained further south (Waldman andBishop unpublished). Additional fieldwork mayreveal a larger range. The recorded locations inNew Zealand were based on aquatic habitatsrather than on frogs found on roads, and this couldexplain why our rate of success in finding L. aureawas higher in New Zealand than in Victoria.

The status of L. aurea in Victoria is unclear.There are very few recorded locations for thisspecies in Victoria based on more than one froghaving been observed, and only one (Swamplandnear Sydenham Inlet) was based on over 100individuals (Gillespie 1996). We found thespecies at only 4 of the 26 previously recordedsites in Victoria that had still, open water whenwe revisited them, and at only one of these sites(Tostaree Pilot Farm) did we detect in excess of10 individuals. Further targeted surveys for thisspecies in Victoria seem warranted.

Gambusia may have had a negative impact on thedistribution of L. aurea in NSW and New Zealand,and could threaten populations of this frog inVictoria. It readily eats the eggs and tadpoles of L.aurea when provided with them in the laboratory,and breeding by this frog species is mostly restrictedto sites where this fish is absent (Morgan andButtermer 1996; Pyke and White 1996, 2000;White and Pyke 1996). Its presence was associatedwith sites in New Zealand where we failed to detectL. aurea but where it had been previously recorded.Gambusia may be absent from L. aurea habitat in



Victoria, but it is abundant in geographicallynearby locations (Pyke and White unpublished).Furthermore, given the exceptionally broad habitattolerances of Gambusia (Weatherley and Lake1967; Arthington and Lloyd 1989; Lloyd et al.1986) and the presence of apparently suitablehabitat in Victoria (Pyke and White, unpublished),there would seem to be considerable potential forfurther spread of this fish into Victorian sitesoccupied by L. aurea.

The observed differences in L. aurea habitat inNSW, Victoria, and New Zealand may reflect therelative abundance of habitat types rather thandifferences in preferred frog habitat. In NewZealand, for example, most recorded sites were ineither garden or farmland areas where waterbodies tended to be relatively small and shallow.Water in these sites was likely to be affected bypollution via runoff from grazing or humanhousehold activities (Tables 4-5). In Victoria,many of the recorded sites are away from urbanareas and hence from areas with high levels ofhabitat disturbance. These sites receive some oftheir water from areas of natural vegetation ratherthan from other sources (Tables 4-5). In NSWmuch of the original L. aurea habitat has beenaffected by urban and industrial expansion andvery little remains untouched (Pyke and White1996, 2001; White and Pyke 1996). Water bodiesin NSW, where L. aurea occurs, are often nearurban areas or areas of bare ground (e.g., quarries,building or industrial sites) (Tables 3 and 5).These areas and studies are therefore combined torevise descriptions of habitat for this species.

Although the present study increases the numberof L. aurea breeding sites that we have examinedfrom 19 to 39, the general habitat description ofbreeding habitat has changed little. By combiningall results (Table 5), it can be seen that breedinghabitat for L. aurea generally contains water bodiesthat are still (i.e., 39/39), small (i.e., 29/39 are<1000 sq. m.), shallow (i.e., 30/39 are <1m),unshaded (i.e., 33/39), free of Gambusia (i.e.,32/39), with emergent aquatic vegetation (i.e.,30/39), and with potential shelter provided bynearby rocks or thick, low vegetation (i.e., 38/39).It is also apparent that most of the known currentbreeding sites for this species are moderately tohighly disturbed by human activities (i.e., 32/39).In addition, ponds where L. aurea breeds are eithersemi-permanent with significantly fluctuatingwater level or are completely ephemeral (i.e.,34/39), rather than being predominantlyephemeral, and have a substrate of sand, rocks orclay (i.e., 34/39), rather than predominantly just

sand or rocks, thus expanding upon what wassuggested earlier (Pyke and White 1996).

Litoria aurea is apparently quite flexible in terms ofboth the general habitats it uses and its aquaticenvironment. While sites where L. aurea breedsusually have the above characteristics, many pondattributes clearly are not mandatory and the speciesis able to breed under a wide variety of habitatconditions. It breeds, for example, at some siteswhere predatory fish are present (introducedGambusia sp. or native Anguilla sp.), at some siteswhich lack emergent aquatic plants, and at somesites where shelter is provided by rocks rather thanthick, low vegetation (Table 5). It is apparentlysimilarly flexible in terms of substrate, with noapparently unsuitable substrate type, and in termsof nearby terrestrial vegetation and the source ofwater at breeding sites (Table 5). The ranges ofphysical and chemical properties that we havefound for water containing L. aurea tadpoles aresimilarly very broad (Table 6).

However, L. aurea appears to restrict its breedinghabitat to water bodies that are still, sunny and lowin salinity. For all 39 L. aurea breeding sites that wehave observed, water has been still, and at none ofthese sites has the water body been largely orcompletely shaded (Table 5). We did not detect thisspecies at sites where the water was either flowingor completely shaded (Table 5). In addition, themaximum observed salinity in water containing L.aurea tadpoles was 7.3 ppt (Table 3 and Table 5)and mass tadpole death has been observed onBroughton Island when salinity has increasedabove this level through intrusion of sea-water intothe ponds there (Pyke and White unpubl.). Sea-water has a salinity of about 34 ppt (Pyke unpubl.).

Litoria aurea might be considered a “weedy” or“colonizing” species. A high level of disturbancehas been found to be a ubiquitous feature of itsbreeding habitat. It also has relatively highfecundity and dispersal ability and is relativelylong-lived (Pyke and White 2001). These traitsshould enhance its ability to survive periods whenconditions are unsuitable for breeding, as well asto colonise and exploit sites when disturbancerenders them more suitable for breeding. Atpresent, however, it is not known how disturbancemight enhance breeding habitat.

Paradoxically, the frogs most often encountered inNew Zealand are all Australian species and, whilefew Australians have ever encountered either L.aurea or L. raniformis, they are the only frogs thatmany New Zealanders have ever seen. The nativefrog species of New Zealand are generally secretiveand have restricted ranges generally far from




Pyke et al.

agricultural or urban landscapes where Australianspecies frequently thrive and where peoplegenerally spend their time (Bell 1982a; Newman1996; Thurley and Bell 1994; Gill and Whitaker1996). Thus there is little spatial overlap betweenthe introduced Australian frog species and theNew Zealand native frog species, and NewZealand native frogs are seldom encountered (Bell1982a; Newman 1996). In Australia L. aurea andL. raniformis are relatively rare, whereas in NewZealand they are locally common and frequentlyencountered species (Pickard and Towns 1988;Ehmann 1996; Tyler 1997).

New Zealand may provide unique opportunities forfurther research into the biology of Australian frogs.That Australian frogs are commonly encounteredin New Zealand suggests that their populationnumbers there may be relatively high. Furthermore,unlike Australia where most frog communitiesinvolve at least two and often many more frogspecies (e.g., Tables 1 and 3; Pyke and White 1996),those in New Zealand usually have just a single

Australian species and occasionally one or twoadditional Australian species (e.g., Table 2; Bell1982b; Pickard and Towns 1988; Pyke and White1996). In addition, the areas surrounding L. aureaand L. raniformis sites in New Zealand are generallyartificial, usually lack native plants, and hence maybe relatively simple in terms of communitystructure. Hence field manipulations of thesespecies may be readily possible without affecting anyNew Zealand native plants or animals. Finally, L.aurea and L. raniformis are presently consideredthreatened species in Australia and have been givena high level of legal protection under both State andFederal law (e.g., Edgar and Stephens 1993; Lunneyand Ayers 1993; Mansergh et al. 1993; White andPyke 1996), whereas they are unprotected feralspecies in New Zealand (Pickard and Towns 1988).Hence, it may be possible in New Zealand to studyrelatively large populations of these Australian frogspecies in relatively simple communities andhabitats that can be experimentally manipulatedwithout legal constraints.

AcknowledgementsThis Project was supported by the AustralianMuseum, the University of Canterbury, the NewZealand Lottery Grants Board Science Research,and the authors’ personal resources. Manyvolunteers helped in the field. Greg Gowing and

Marie Gillon have helped with computerisation ofthe data. Helpful comments on an earlier draft ofthis paper were provided by Ross Goldingay, BradLaw, Dan Lunney and an anonymous referee. Forall this assistance we are most grateful.

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Table 1. Sites visited in Victoria. All locations are in AMG Zone 55.

Site Name

SandyWaterholeCreek

Nash Camp

Wallagaraugh

Tobins Creek

Genoa

Cape ConranQuarry

West Yeerung

Cape Conran

Point Ricardo

Wongabeena

Marlo Plains

Scrubby Creek

Genoa PeakRoad #1

Genoa PeakRoad #2

MallacootaAirport

Burglar Gap

Three Sisters

Pettmans Beach

Morass Break

Hospital Creek

Old OrbostTrack

Map GridReference

E7297N58589

E7279N58536

E7382N58498

E7254N58479

E7296N58486

E6523N58173

E6562N58243

E6527N58589

E6423N58147

E6441N58154

E6451N58164

E7281N58482

E7279N58476

E7299N58449

E7403N58351

E7182N58576

E6832N58615

E6044N58123

E6026N58130

E6079N58162

E6024N58158

Elevation(m)

160

130

0

40

40

20

90

15

35

35

35

10

20

240

25

120

690

5

30

5

60

PreviousL. aureaRecords

1981

1991/92

1991/92

1968

1990

1990 and1992

1990/91

1990/91and

1990/91

1990/91

1990/91

1988/89

1988/89

1988/89

-

1984/85

1995/96

1992/93

1992/93

1992/93

1992/93

Present L. aureaObservations

Tadpoles

Five males calling;1 female captured

Other Frog SpeciesDetected

Crinia signifera

Litoria phyllochroa

Litoria ewingii

Litoria peroniiCrinia signifera

Crinia signifera

Crinia signiferaLitoria peronii

Nature ofWater BodiesPresent

Slow flow

Still

Still

None

Still

Still

Moderate flow

Still

None

Still

Still

Still

Moderate flow

None

Still

None

Still

Still

Still

Still

None

GambusiaPresent?

No

No

No

No

No

No

No

No

No

No

No

No

No

No

No

No

No

No

No

No

No




Pyke et al.

Tostaree PilotFarm

Montas Break

BonangHighway

Tostaree

Lake Beatle

Little LakeBeatle

Ewing Marsh

GippslandRailway

Simpsons CreekRoad

Princes Hwy #1

Princes Hwy #2

Princes Hwy #3

Wombat Track

Thurra River

Reedy Creek

Thurra Road

Gibbs Creek

Tamboon

Tamboon River

Bobs Bay

Bemm River

Pearl Point

E6046N58173

E6047N58161

E6303N58308

E6035N58211

E6246N58178

E6240N58169

E6235N58126

E6179N58208

E6172N58206

E6071N58207

E6152N58203

E6142N58205

E6132N58204

E6026N58158

E6936N58368

E6904N58283

E6912N58244

E6899N58205

E6882N58227

E6734N58201

E6719N58201

E6705N58170

50

60

60

35

50

25

5

15

40

55

50

25

40

20

200

110

95

140

5

2

5

5

1992/93

1992/93

1991/92

1992/93

1992/93

1992/93

1992/93

1992/93

1992/93

1992/93

1992/93

1992/93

1992/93

Nonerecorded

1991/92

1991/92

1991/92

1991/92

1991/92

1990/91

1990/91

1990/91

At least 30 adultsobserved; malescalling; tadpolesobserved

Adults observedand Males calling

Crinia signiferaLimnodynastesdumerilii

Paracrinia haswelliLitoria peroniiLimnodynastes dumeriliiLitoria ewingii

Crinia signifera

Crinia signiferaLimnodynastesdumeriliiLitoria ewingii

Litoria ewingiiLimnodynastesdumeriliiUperoleia tyleri

Litoria lesueuri

Litoria phyllochroa

Litoria phyllochroa

Crinia signifera

Still

Still

None

Still

Still

Still

Still

None

None

None

None

None

None

Still

Moderate flow

None

Moderate flow

None

Still

Still

None

Still

No

No

No

No

No

No

No

No

No

No

No

No



Site

Nam

e

Taur

aroa

Waio

tira

#’s

1,2

and

3

Gle

nber

vie F

ores

t

Lake

Waip

oroh

ita

Toke

rau

Beac

h

Flet

cher

Bay

#1

Flet

cher

Bay

#2

Port

Jack

son

Opo

tiki

Waio

eka

Rive

r

Kape

nga

Swam

p

Roto

rua

Map

Grid

Refe

renc

e

E262

0,N

6588

8

E261

8,N

6585

5

E263

0,N

6621

8

E264

3,N

6700

0

E254

5,N

6695

3

E272

5,N

6521

4

E272

5,N

6521

8

E272

0,N

6521

3

E289

3,N

6342

9

E288

7,N

6324

5

E279

3,N

6324

4

E279

4,N

6339

2

Elev

atio

n (m

)

60 100

220

10 10 10 10 10 10 10 360

270

Prev

ious

L.a

urea

Rec

ords

L.au

rea

reco

rded

in 1

991/

92

L.au

rea

reco

rded

in 1

991/

92

and

earli

er a

t #

’s 1

and

2

L.au

rea

reco

rded

in 1

992

L.au

rea

reco

rded

in 1

985/

86

L.au

rea

reco

rded

and

dat

eun

know

n

L.au

rea

reco

rded

in 1

989

L.au

rea

reco

rded

in 1

989

L.au

rea

reco

rded

in 1

989

L.au

rea

reco

rded

in 1

995/

96

L.au

rea

reco

rded

and

dat

eun

know

n

L.au

rea

reco

rded

in 1

982

L.au

rea

reco

rded

and

dat

eun

know

n

Pres

ent

Obs

erva

tions

ofL

.aur

eaan

d O

ther

Fro

g Sp

ecie

s

No

tadp

oles

or

frogs

No

tadp

oles

or

frogs

at

#’s

1 an

d 2;

tadp

oles

of L

.aur

eaan

dL.

ewin

giia

t #

3

Tadp

oles

,met

amor

phs

and

imm

atur

es

of L

.aur

ea

No

tadp

oles

or

frogs

Frog

s of

L.au

rea

and

L.ra

nifo

rmis

obse

rved

;11

L.a

urea

cap

ture

d,in

clud

ing

7 m

ales

inpr

esen

t or

pos

t-re

prod

uctiv

e co

nditi

on a

nd4

mat

ure

fem

ales

L.au

rea

tadp

oles

.

Tadp

oles

and

frog

s of

L.a

urea

.

L.au

rea

frogs

No

tadp

oles

or

frogs

No

tadp

oles

or

frogs

No

tadp

oles

or

frogs

No

tadp

oles

or

frogs

Com

men

ts r

e G

ambu

sia

Gam

busia

abun

dant

Gam

busia

abun

dant

at #

1 an

d no

Gam

busia

reco

rded

at #

’s 2

and

3

No

Gam

busia

reco

rded

Gam

busia

abun

dant

No

Gam

busia

reco

rded

No

Gam

busia

reco

rded

No

Gam

busia

reco

rded

No

Gam

busia

reco

rded

No

Gam

busia

reco

rded

No

area

of s

till o

r slo

w w

ater

No

Gam

busia

reco

rded

No

Gam

busia

reco

rded

Table 2: Sites visited in New Zealand




Pyke et al.

Site

Nam

e

Des

crip

tion

Map

Grid

Refe

renc

e

Elev

atio

n

Surv

ey D

ates

Reco

rd

Subs

trat

e

Wat

er S

ourc

e

Wat

er A

rea

(sq.

m.)

Max

imum

Dep

th (c

m)

Rate

of

Wat

er F

low

Wat

er L

evel

Shad

e

Emer

gent

Aqu

atic

Vege

tatio

n

Mer

oo L

ake

Nat

ural

coas

tal

wet

land

E262

9N

6070

2

5 Nov

’99

&

Feb

‘00

Call

ing

male

s &

tadp

oles

Sand

Vege

tatio

n

60 35 Still

E Part

ial

Junc

us s

p.

Cur

rum

bene

Disu

sed

clay

quar

ry

E287

8N

6129

0

40 Sept

‘99

Call

ing

male

s

Clay

Vege

tatio

n

150

200

Still

F Non

e

Typh

a sp

.El

eoch

aris

sp.

Mer

rylan

ds

Disu

sed

clay

quar

ry

E314

3N

6254

5

40 Jan

‘00

Tadp

oles

Clay

Vege

tatio

nBa

re G

roun

d

40 15 Still

E Non

e

Typh

a sp

.Ju

ncus

sp.

Lake

Con

jola

–Bu

ckle

ys P

oint

Serie

s of

natu

ral d

une

swale

pon

ds

E270

7N

6091

9

5 Jan

‘00

Call

ing

male

s &

tadp

oles

Sand

Vege

tatio

n

2000

55 Still

E Part

ial

Scho

enop

lect

ussp

.

Lake

Con

jola

– Pa

ttim

ores

Lago

on

Art

ificial

pond

age

crea

ted

byem

bank

men

t

E270

9N

6092

5

20 Feb

‘98

Call

ing

male

s

Sand

Vege

tatio

n

120

45 Still

F Non

e

Typh

a sp

.Ju

ncus

sp.

Brou

ghto

nIsl

and

pond

s

Nat

ural

pond

sad

jacen

t to

ocea

n

E436

4N

6391

1

5 Mar

ch ’9

8 –

Mar

ch ‘0

0

Call

ing

male

s &

tadp

oles

Rock

Vege

tatio

n

0.1-

25

45 Still

E Non

e

Mos

tly n

one

& o

ne h

asPh

ragm

ites

sp.

Nor

th A

voca

Art

ificial

pond

age

crea

ted

by e

mba

nkm

ent

adjac

ent t

oco

asta

l lago

on

E354

2N

6296

1

5 Nov

-Dec

‘98

Call

ing

male

s &

tadp

oles

Sand

Vege

tatio

nU

rban

1600

70 Still

E Part

ial

Phra

gmite

s sp

.

Dur

ras

Nor

th

Dam

E255

4N

6052

7

15 Dec

‘96

Call

ing

male

s &

tadp

oles

Sand

Vege

tatio

n

2800

~40

0

Still

F Non

e

Non

e

Pt.K

embl

aSt

eelw

orks

Art

ificial

pon

d

E306

2N

6183

1

10 Dec

‘96

Call

ing

male

s &

tadp

oles

Con

cret

e

Indu

stria

lBa

re g

roun

d

400

>1m

Still

F Non

e

Typh

a sp

.Cy

peru

s sp

.

Coo

mad

itchy

Sem

i -na

tura

lw

etlan

dsu

rrou

nded

by

park

land

&su

burb

ia

E307

0N

6181

4

20 Jan

‘96

Call

ing

male

s &

tadp

oles

Sand

Park

land

Urb

an

15,0

00

>2m

Still

F Non

e

Typh

a sp

.El

eoch

aris

sp.

Mar

rickv

ille

Art

ificial

pon

d in

small

sub

urba

nnu

rser

y

E329

9N

6246

9

20 Feb

’99

–Fe

b ‘0

0

Call

ing

male

s &

tadp

oles

Clay

Tow

n

50 60 Still

F Part

ial

Typh

a sp

.

Kurn

ell

* * * Oct

’98

-Jan

‘00

Call

ing

male

s&

tadp

oles

* * * * * * * *

Tabl

e 3.

Hab

itat

info

rmat

ion

for

L.au

rea

bree

ding

site

s in

NSW

(Se

e Py

ke a

nd W

hite

199

6 fo

r ha

bita

t in

form

atio

n fo

r Ku

rnel

l Site

) an

d ph

ysic

al/c

hem

ical

pro

pert

ies

of w

ater

cont

aini

ng L

.aur

eata

dpol

es.A

ll lo

catio

ns a

re in

AM

G Z

one

56.



Terr

estr

ial

Vege

tatio

nw

ithin

50m

Pollu

tion

Pote

ntia

lSh

elte

r

Hum

anD

istur

banc

e

Oth

er F

rog

Spec

ies

Pred

ator

y Fi

sh

Turb

idity

(nts

)

Diss

olve

dO

xyge

n (g

/l)

ORP

(mvo

lts)

pH Salin

ity (p

pt)

Wat

er T

emp

(deg

.C)

Woo

dlan

d

Non

e

Vege

tatio

n

Non

e

Litor

ia d

enta

taLim

nody

nast

espe

roni

iCr

inia

sign

ifera

Angu

illa s

p.

2-52

4.4-

12.8

41-1

19

6.7-

7.7

0.1-

0.9

16-2

2

Fore

stSh

rubl

and

Bare

Gro

und

Non

e

Vege

tatio

n

High

Litor

ia p

eron

iiLim

nody

nast

espe

roni

i

Non

e

Shru

blan

dBa

re G

roun

d

Non

e

Vege

tatio

nRo

cks

Tim

ber

Art

ificial

High

Litor

ia p

eron

iiLim

nody

nast

es

pero

nii

Crin

ia s

ignife

ra

Non

e

6 4.2

117

6.5

0.1

21

Fore

st

Non

e

Vege

tatio

n

Non

e

Limno

dyna

stes

pero

nii

Crin

ia s

ignife

raPs

eudo

phry

ne

bibr

oni

Non

e

Gra

sslan

d

Gar

bage

Vege

tatio

nA

rtific

ial

High

Limno

dyna

stes

pero

nii

Crin

ia s

ignife

ra

Non

e

Shru

blan

dBa

re G

roun

d

Non

e

Vege

tatio

n

Non

e

Limno

dyna

stes

pero

nii

Non

e

0-14

6

3.3-

16.0

-531

-351

5.7-

9.8

0.2-

7.3

19-3

7

Gra

sslan

dW

oodl

and

Gar

den

Urb

an

Vege

tatio

nA

rtific

ial

High

Litor

ia p

eron

iiL.

tyler

i L.d

enta

taL.

caer

ulea

Limno

dyna

stes

pero

nii

Non

e

19-3

1

3.3-

9.1

143-

192

5.7-

6.7

3.1-

6.0

22-2

5

Gar

den

Non

e

Vege

tatio

nA

rtific

ial

High

Litor

ia p

eron

iiLim

nody

naste

spe

ronii

Crin

iasig

nifer

aU

pero

leia

laev

igata

Non

e

1 7.2

75 7.1

0.7

22

Bare

gro

und

Indu

stria

l &

Gar

bage

Vege

tatio

nRo

cks

High

Non

ere

cord

ed

Non

e

3 3.8

49 8.3

0.1

23

Gra

sslan

d

Gar

bage

Vege

tatio

n

Med

ium

Non

e re

cord

ed

Gam

busia

17 1.0

77 8.2

0.1

24

Bare

Gro

und

Gar

den

Non

e

Vege

tatio

n

High

Limno

dyna

stes

pero

nii

Non

e

34-8

9

4.0-

10.5

32-1

56

6.0-

7.6

0.1-

0.4

14-2

7

* * * * * * 2-14

8

3.4-

14.6

-11-

148

5.4-

9.1

0.0-

1.2

12-3

3




Pyke et al.

REG

ION

Site

Nam

e

Des

crip

tion

Subs

trat

e

Wat

er S

ourc

e

Wat

er A

rea

(sq.

m.)

Max

imum

Dep

th (c

m)

Rate

of W

ater

Flo

w

Wat

er L

evel

Shad

e

Emer

gent

Aqu

atic

Vege

tatio

n

Terr

estr

ial Ve

geta

tion

with

in 5

0m

Pollu

tion

Pote

ntial

She

lter

Dist

urba

nce

Pred

ator

y fis

h

Turb

idity

(nts

)

Diss

olve

d O

xyge

n (g

/l)

Oxi

datio

n Re

duct

ion

Pote

ntial

(mvo

lts)

pH Salin

ity (p

pt)

Wat

er Te

mpe

ratu

re

(deg

.C.)

Cap

e C

onra

nQ

uarr

y

Disu

sed

rock

quar

ry

Rock

Vege

tatio

nBa

reG

roun

d

900

30 Still

E Non

e

Scho

enop

lectu

s sp

.Cy

peru

s sp

.

Fore

st a

nd B

are

Gro

und

Non

e

Vege

tatio

n

Hig

h

Non

e

27.5

7.0

275

4.4

0 31

Tost

aree

Pilo

tFa

rm

Dam

Cla

y

Gra

zing

3,00

0

>1m

Still

F Non

e

Eleo

char

is sp

.

Gra

ssla

nd

Gra

zing

Vege

tatio

n

Hig

h

Non

e

4.5

7.1

105

6.6

0.2

23

Thur

ra R

iver

Nat

ural

wet

land

adjac

ent t

o riv

er

Sand

Vege

tatio

n

15,0

00

40 Still

F Non

e

Phra

gmite

s sp

.Cy

peru

s sp

.

Fore

st

Non

e

Vege

tatio

n

Non

e

Non

e

Mal

laco

ota

Airp

ort

Dam

Sand

Park

land

2,80

0

>1m

Still

F Non

e

Eleo

char

is sp

.

Fore

st

Non

e

Vege

tatio

n

Hig

h

Non

e

Wai

otira

#3

Dam

Cla

y

Gra

zing

800

>1m

Still

F Non

e

Azol

la s

p.Ju

ncus

sp.

Alism

a sp

.

Gra

ssla

nd

Gra

zing

Vege

tatio

n

Hig

h

Non

e

17.6

8.7

86 7.8

0 26

Gle

nber

vie

Fore

st

Dam

Cla

y

Vege

tatio

n

10,0

00

>7m

Still

C Non

e

Eleo

char

is sp

.

Fore

st a

ndSh

rubl

and

Non

e

Vege

tatio

n

Hig

h

Non

e

3.6

6.7

112

5.7

0 23

Toke

rau

Beac

h

Dam

Sand

Gra

zing

200

50 Still

F Non

e

Non

e

Gra

ssla

nd

Gra

zing

Vege

tatio

n

Hig

h

Non

e

Flet

cher

Bay

#1

Cat

tle w

ater

trou

gh

Con

cret

e

Vege

tatio

n

1 40 Still

F Non

e

Non

e

Gra

ssla

nd

Gra

zing

Vege

tatio

n

Hig

h

Non

e

103.

2

9.1

152

9.5

0 22

Flet

cher

Bay

#2

Smal

l pon

dage

on c

lear

edgr

azin

g la

nd

Sand

Gra

zing

100

40 Still

F Non

e

Cype

rus

sp.

Gra

ssla

nd

Gra

zing

Vege

tatio

n

Hig

h

Non

e

119.

3

5.4

65 6.4

0 24

VIC

TOR

IAV

ICTO

RIA

Tabl

e 4.

Hab

itat

info

rmat

ion

and

phys

ical

/che

mic

al p

rope

rtie

s of

wat

er fo

r L.

aure

abr

eedi

ng s

ites

in V

icto

ria a

nd N

ew Z

eala

nd



Parameter

Natural vs. artificial

Human disturbance

Substrate

Shade

Water Level

Water Flow

Water Source

Pollution

Max.Water Depth(cm)

Water Surface Area(sq.m.)

Nearby TerrestrialVegetation

Category

Mostly or completelynatural

Largely or entirelyhuman-created

High

Medium

None

Sand

Clay

Rock

Concrete

Rubble

None

Partial

Complete

Ephemeral

Fluctuating

Constant

Still

Slow

Rapid

Bare Ground

Parkland

Grazing

Natural Vegetation

Urban

Industrial

None

Likely

Grazing

Urban

Industrial

Garbage

0-50

50-100

>100

0-100

101-1000

>1000

Bare Ground

Garden

Grassland G

NSW (fromPyke andWhite 1996)

2

17

8

8

3

10

4

1

2

2

17

2

12

3

4

19

8

5

4

2

7

1

8

11

3

8

1

11

7

1

9

8

2

8

6

9

NSW (this study)

3

8

7

1

3

6

5

7

4

5

6

11

2

1

8

2

1

7

4

1

1

3

4

3

4

4

3

4

5

3

2

Victoria

1

3

3

1

2

1

1

4

1

3

4

1

1

1

2

3

1

1

2

2

1

3

1

1

New Zealand

0

5

5

2

2

1

5

4

1

5

3

2

1

4

4

3

2

2

2

1

4

NSW,Victoriaand NewZealand (this study)

4

16

15

1

4

10

9

1

16

4

6

13

1

20

3

2

4

12

2

1

11

9

5

1

1

3

9

3

8

6

6

8

6

3

7

Table 5. Summaries of habitat characteristics for breeding sites of L. aurea in NSW,Victoria and New Zealand.Values are numbers of sites.




Pyke et al.

Emergent AquaticPlants

Potential Shelter

Predatory Fish

Heathland

Shrubland

Woodland

Forest

Present

Absent

Typha sp.

Eleocharis sp.

Juncus sp.

Phragmites sp.

Schoenoplectus sp.

Cyperus sp.

Low Vegetation

Rocks

Timber

Artificial

Not recorded

Recorded

Gambusia recorded

Anguilla recorded

1

6

3

14

5

11

4

2

1

16

3

4

14

5

5

3

2

2

9

2

5

1

3

1

1

10

2

1

4

9

2

1

1

3

4

2

1

1

2

4

4

1

1

3

2

1

1

1

5

5

4

2

6

16

4

5

4

4

1

1

4

19

2

1

4

18

2

1

1

Region NSW Victoria New Zealand

Turbidity (nts) 0-148 5-28 4-119

Dissolved Oxygen (g/l) 3-16 7 5-9

ORP (millivolts) -531 – 351 105-275 65-152

pH 5.4-9.8 3.7-6.6 5.7-9.5

Salinity (ppt) 0.1-7.3 0-0.2 0

Water Temperature (Deg. C) 12-37 23-31 22-26

Number of sites 8 2 4

Table 6:Comparison of physical/chemical properties of water containing L. aurea tadpoles in NSW,Victoria andNew Zealand.

Appendix: Description of how location records for L. aurea wereobtained and selected.

We obtained information about the known L. aurea sites in Victoria from avariety of sources. In August 1996 the Victorian Department of NaturalResources and Environment provided us with a copy of all records of L. aurea inthe Atlas of Victorian Wildlife.This yielded 120 records, representing 94 distinctsites. In November 1996 we obtained location information for L. aureaspecimens held by both the Australian Museum and the Victorian Museum.Thisyielded 3 records from 2 sites for the Australian Museum collection and 73records from 26 sites for the Victorian Museum collection.All but one of theseMuseum records were included in the Atlas of Victorian Wildlife data, and thisone record was from a suburb of Melbourne and was considered to be probablyan animal that had escaped from captivity.During and before our trip to Victoriawe learnt about a further three recorded sites via personal communication. Forour fieldtrip in Victoria we therefore targeted the sites in the Atlas of VictorianWildlife plus these three additional sites, for a total of 97 sites.

AP

PE

ND

IX



We obtained information about known L. aurea sites in New Zealand fromsimilar sources. In late 1996 the New Zealand Department of Conservationprovided us with all the reliable L. aurea records contained in its Amphibianand Reptile Distribution Database. This resulted in 31 distinct site recordscovering the period 1982-1993.Although this database, which was originallyknown as Biosite, had been maintained since 1951, it relied heavily onvolunteers who did not distinguish L. aurea and L. raniformis before 1982 (Bell1982a,b) and from 1993 until 1999 the database was poorly maintained withno L. aurea records being added during this period. All of the records werefrom the North Island, except for a single record from the northern part ofthe South Island, which may have been the result of a misidentification as asurvey of the site in 1997 yielded specimens only of Litoria ewingii (Bishop andWaldman unpublished). In 1996, following press releases asking forinformation regarding current frog distributions, many letters were receivedcontaining descriptions of frog observations. By the end of 1996 this hadgenerated an additional 15 recorded L. aurea sites, of which we consideredthree to be reliable.We located and surveyed 41 of the 97 targeted sites in Victoria (Table 1).Weattempted to locate 61 of the 75 sites east of Lake Tyers, with timeconstraints preventing us from surveying 22 sites that were further west andfrom surveying 14 sites that were less readily accessible.There were also 20sites that we could not confidently find, because their site coordinates wererecorded only by latitude and longitude in degrees and minutes and wereclearly approximations, as evidenced by the fact that some of them fell withinseawater at some distance from dry land. We were able to find locationswhere Australian Map Grid reference coordinates were available.For the purposes of the present study we chose 11 L. aurea sites to visit inNew Zealand. This we did by first plotting all the known sites on 1:50,000maps, then highlighting those sites where the largest numbers of L. aurea hadbeen reported and finally choosing a route that would include sites coveringmost of the known range of L. aurea and as many as possible of the locationsin the available time.We located and surveyed all eleven sites (Table 2).

AP

PE

ND

IX


Habitat-use by the Green and Golden Bell Frog Litoria aurea in Australia and New Zealand

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Transcript of Habitat-use by the Green and Golden Bell Frog Litoria aurea in Australia and New Zealand