Devonian palaeobiogeography of Australia and adjoining regions

4

Devonian palaeobiogeography of Australia and adjoining regions

J.A. TALENTl, R. MAWSON!, J.C. AITCHISON', R.T. BECKER', KN. BELLI, M.A. BRADSHAW., C,J. BURROW', A.G. COOK', G.M. DARGAN',

J.G. DOUGLAS', G.D. EDGECOMBE', M. FEISTIO P.J. JONESlI, J.A. LONG12, J.R. PHILLIPS-ROSS13, J.w. PICKETT', G. PLAYFORDl., R.B. RICKARDSl',

B.D. WEBBYl, T. WINCHESTER-SEETOI, A.J. WRIGHTl', G.c. YOUNGlI & Y._Y.ZHEN9

lCentre for Ecostratigraphy and Palaeobiolog)~ Department a/Earth and Planetmy Sciences, Alacquarie University 21 09, NSTf~ Australia

2Department a/Geology, University a/Hong Kong, Pokfidam Rd, Hong Kong, China 3Museumjiir Naturkunde, Humboldt-Universitiit, Inl'alidenstrasse 43, D-I0155 Berlin, Germany 4Geo!ogica[ Sciences, University ofCanterbur)~ Private Bag 4800, Christchurch. New Zealand 5Department a/Zoology, University a/Queensland, Brisbane 4072, Queensland,Australia 6Queensland Afl/seum, P. 0. Box 3300, South Brisbane 4101, Queensland, Australia 7 Geological Survey ofNSW, P.O. Box 76, Lidcombe 2141, NSW; Australia 841 Grieve Street, Warrnambool, VictOJ'ia 3280, Australia 9 Australian Museum, PO Box A285, Sydney South 2000, NSW, Australia 10 Laboratoire de Paleobotanique, Universite. deMontpellier-2, Place Bataillon, 34095 Alontpelliel; France 11 Department ofGeolog)~ Australian National Universit)~ Canberra 0200, ACT, Australia 12/Vestel71 Auso-alian Museum, Francis St, Perth 6000, WA, Australia 13Department of Biology, Western Washington University, Bellingham, WA 98225, USA 14DepartmentofEarth Sciences, University afQueensland, Brisbane 4072, Queensland, AlIso'alia 15 Department of Earth Sciences, University afCambridge, Downing St, Cambridge CB2 3EQ, u.K. 16Schaal afGeosciences, Unil'ersityofWollongong, Wol!ongong 2522, NSW; Australia

PALAEOBIOGEOGRAPHIC analyses are only as good as their chronologic underpinning. Zonal schemes for substantial parts of Devonian time have been proposed based on graptolites (Early Devonian: Jaeger, 1988), daclyoconarids (Early and Middle Devonian: Albelti: 1987-2000), ammonoids (Emsian to Famennian: House, 1973), chitinozoans (Lochkovian-Frasnian: Paris, 1981)

and spores (Richardson & McGregor, 1986). Zonal schemes based on conodonts, developed over the past 30 years, embrace all of Devonian time (Klapper & Johnson, 1980; Sandberg & Dreesen, 1984; Klapper, 1989; Ziegler & Sandberg, 1990; Klapper & Becker, 1999). Inevitably, modifications of all these schemes continue to be proposed (e.g., Yolkin et al .• 1994;

TALENT, JA, MAWSON, R.,AITCHISON, J.C., BECKER, R.T., BELL, K.N., BRADSHAW, M.A., BURRo\V; C.J., COOK, A.G., DARGAN, G.M., DOUGLAS, lG., EDGECOMBE, G.D., FEIST, M., JONES, P.I, LONG, I.A., PHILLIPS-ROSS, J.R., PICKETT, J.W., PLAYFORD, G., RICKARDS, R.B., WEBBY, B.D., WINCHESTER-SEETO, T., WRIGHT, AJ., YOUNG, G.C. & ZHEN, Y.-Y., 2000:12:20. Devonian palaeobiogeography of Australia and adjoining regions, Memoir of the Association of Australasian Poloeontalagists 23, 167-257. ISSN 0810-8889

168

Early Devonian (ca. 400 Ma)

Late Devonian (ca. 365 Ma)

Early Devonian

Fig. 1. Location of Australia and other cmstal blocks surrounding Prototethys for two intervals: Early Devonian, around 400 Ma, and latc Devonian, around 365 Ma (based on Li & Powell, 2000). In the Devonian Australia stayed in the tropics and the adjoining Gondwana supercontinent rotated anticlockwise during the interval between reconstructions depicted here. Brachiopod data given elsewhere (Talent et al., 2000; Yolkin et al., 2000) and discussed herein, show an almost 100% provincial contrast at species-level during Early Devonian and Eifelian times between Australia and the South China Block (SCB), and a linkage between faunas of the North China Block (NCB) and Europe (Baltica). These are consistent with both blocks being located farther away from Australia during Early and Middle Devonian times, and with North China being located \V of South China. Brachiopod data, though consistent with this configuration, do not constrain the location of the above blocks. Abbreviations are: BaltBaltica; elM - Cimmerian collage; KAZ - Kazakhstan collage; Sib - Siberian continent.

AAP Memoir 23 (2000)

Valenzuela-Rios & Murphy, 1997, 1999) for conodonts for portions of the Early Devonian. Applicability of these proposals-particularly constraints imposed by provinciality of the pivotal organisms-have yet to be tested, even semiglobally.

Conodont data obtained from E Australia, mainly over the past IS years, have resulted in significant stratigraphic re-alignments for rocks of Devonian age and better understanding of the transgression-regression pattern (Mawson & Talent, 2000). Recent surveys of the Late Devonian ofthe Canning Basin ofNW Australia, based on conodonts and cephalopods (Becker & House, 1997; Becker ef al., 1993), supplemented by the synthesis for the western Australian sedimentmy basins given by MOlY & Beere (1988), provide adequate coverage for Western Australia. The framework provided by these syntheses forms the basis for the analysis of Devonian biogeography as well as palaeogeography for Australia and adjoining regions presented below (Figs. 1-4)1. Because of the potential for readers to become lost in a plethora of stratigraphic names for the Australian sequences mentioned in this discussion, we have included a table of stratigraphic alignments as Appendix 2.

Brachiopods have been pivotal in discussions on Devonian marine biogeography. The significance of the Early Devonian-EifeHan brachiopods has been evaluated qualitatively on several occasions, principally about 30 years ago (Boucot ef al., 1967, 1969, 1975; Talent, 1972), and subsequently more quantitatively for Early Devonian brachiopods (Savage ef al., 1979), each time at generic level. These analyses led to a broad generalisation: that during Early Devonian times the brachiopod faunas, viewed globally, were provincial, and, in the case of the Australian and New Zealand brachiopod faunas, could be regarded usefully as constituting a Tasman Province-with provinciality increasing through Lochkovian to early Emsian times and declining thereafter. This evaluation proved fruitful as a "backdrop" against which to evaluate-always qualitatively-the potential palaeobiogeographic relationships of other groups, notably rugose corals, trilobites, conodonts and chitinozoans (Zhen ef al., 2000; Campbell & Davoren, 1972; Telford in Talent, 1972; Telford, 1979; Talent ef al., 2000; Winchester-Seeto, 1997). There have been no palaeobiogeographic analyses, qualitative or quantitative, of post-EifeHan Devonian

Fig. 2 (opposite). Eastern Australian Early and Middle Devonian localities referred to in the text. Stratigraphic columns for the principal area are given in Appendix 2.


o I

t N

I 500km

I

I Waratah

Bay

'Fanning Downs' Charters Towers

Ukalunda

Yarrol Forearc Complex

Sulcar Attunga Moore Ck

169

Nemingha Tamworth Pigna Barney Mudgee Stuart Town

___ Rylstone Cumnock Euchareena Limekilns Orange

Buchan Tabberabbera Walhalla Coopers Creek Toongabbie Tyers

Bungonla Goulburn Windellama Yass Taemas Wee Jasper Canberra

170

o t

(aproximately rhenana Zone) transgression

500km , 1


Hodgkinson Basin

Drummond Basin

Fig. 3. Australian Late Devonian sedimentary basins with a selection oflocalities referred to in the text (Fig. 4 provides detail for eastem Australia), and New Zealand Early Devonian terranes with location of the two areas from which Early Devonian faunas have been described. .

t I

_ Takaka Terrane

~ Buller Terrane

Takaka Terrane

I~.:~-Ilntrusives

o 100km t t

Fig. 4 (opposite). Eastern Australian Late Devonian localities referred to in the text.


t N

I

~--~ Approximate limit of late Frasnian ~--~" (approximately rhenalla Zone)

transgression

Hodgkinson "Basin"

Tamworth

----Grenfell '----- Canberra

c-____ Ettrema

======~ Merimbula Eden

Genoa Genoa R. Combienbar Tabberabbera

o 500km

171

172

, I

Pandanus Ck - Broken River


Distribution of Early and Middle Devonian Rocks in Eastern Australia

(exclusive of intrusives) with references to conodont faunas.

Camel Creek (Sloan et af. 1995)

(Telford 1975; Mawson 1987b; Mawson ~ e/ aL 1985; Mawson & Talent 1989; ----'1},) Townsville

, Sloane/af. (995) f1, ~~ River

Burdekin Basin / 0 Charters (Talent & Mawson 1994; II Mt Etna (0 1970

(Brock & Talent 1993) Mawson & Talent unpub) Mawson & TaJenf (997) ~ row.

Douglas Creek (Mawson unpub) ~ b Mt Morgan (Taube ef aJ. unpub)

~ Alpha ~~ Clermont ~oko R.nge, (Mawson in ~ 6.,. ~ R . ampton __ Mt Holly Beds (DnJC(J 1970, " H,nd,-on,' ". '995' • , Craven ,..,« 'J ~ Gladstone Mawson ef al-, 1995; Mawson &

'

Peak Beds ~ Talent unpub.)

, (non m'rine) g ~ f Monlo (Fotdh,m unpub.)

___ J _____ , It} {%.... --- --- NogoaAnticline ~ >;;Q~ (Fordham 1976)

, li.!i.'.·.'.i .•.•• ·.•1."tt" Timl>ury Hills , ")'";} V· Basm n . b

Adavale Basin Yi:!'Hi\ (subsurface) 1'1S ane I (subsurface) (~Mt i; ~ _ Silverwood (Telford 1972; I ~t\r'''''''''''~ tI~ __ , Mawsonunpub.) , ___________ w-./" '1,.J Yarramanbully(Malvsonetal.1988;

'

Wellington' Mawson & Talent unpub.} Cobar

J (Pickett 198O) (l1Iilson 1989) Moore Creek (Philip 1987) I - Attunga (Mawson & Talent 1994)

Waratah Bay (BIschoff & AIgenf 1990,

Mawson et af 1992)

Sulcor _ Tamworth (Mawson et al. 1997)

~--f--- Loomberah (Furey-Grieg 1995; Mawson etaf. 1995)

""'1--- Pigna Barney (Dongo/1995) '-;,1'---- Timor (Pedderet al. 1970) r~ __ Mudgee (Pickett1978; Colquhoun 1995)

Nubrigyn (Mawson & Talent 1999)

r--"--....... -- Manildra (Savage 1973; Trotlerunpub.)

,'--''=:r==::::::-- Windellama (MaLvson 1986)

Yass (Unk & Droce 1972)

~---t----- Ravine (Biasulll unpub.)

&1>'00---.1---- Bind! (Mawson 1987a; Mawson ef aI. 1992)

·fY'>:-r---- The Basin (Mawson et al.I992)

X"':"'\-<-.~--- Boulder Flat (MaLvson et aI. 1992) '------ Buchan (Mawson 1987a; Mawson ef aI.1992)

~ ~

Tabberabbera (Mmvson et al. 1992)

'------ Loyola (Cooper 1973; Mawson et aI.1992)

'------Tyers (MaLvson & Talent 1994)

Zeehan .. Queenstown ..

Point Hibbs l t N UM.thinna

(Carey & Berry 1988) Hobart

I


brachiopods, though there has been a widespread assumption that Middle and Late Devonian brachiopod faunas displayed a high level of cosmopolitanism, at least at generic level.

It is worth recalling in this regard that neontologists, when assessing realms, regions and provinces, place greater emphasis on speciessome opting for 50% species contrast in defining provinces, some for less-than palaeontologists who emphasize genera and even higher categories when attempting to define biogeographic entities in the past (cf. Kauffman, 1973; Talent, 1985a). There is thus rarely any sort of coordination between the provinces ofthe neontologist and the provinces of the palaeontologist. In the discussion which follows most emphasis is placed on genera, but an approach to species-level biogeography has been attempted for Devonian conodonts (Flessa & Hardy, 1988; Klapper, 1995; Mawson, herein) and is being attempted for Early and Middle Devonian brachiopods (Talent et ai., 2000; Yolkin et ai., 2000).

Because of their paramount position in establishing intercontinental stratigrapbic alignments, the provinciality of Devonian conodonts, graptolites, chitinozoans, and brachiopods will be considered first, followed by an overview of provincialism with respect to Devonian Brachiopoda in Australia and nearby regions, drawing on the recently revised taxonomy of Early and Middle Devonian brachiopods from the Asia-Australia hemisphere (Talent et ai., 2000, for which the palaeogeographic basis is Talent et ai., 1987). Other marine groups will then be discussed, commencing with the Algae, followed by the ~nonnal' classificatory sequence of marine inveliebrate and veliebrate groups, ending with the non-marine groups, especially plants.

We repeat that, because of the potential for readers to become lost in a plethora of stratigraphic names for Australian Devonian sequences, we have included a table of stratigraphic alignments (Appendix 2) at the end ofthis paper.

MARINE BIOTA Conodonta (R. Mawson)

Pelagic fossil groups, including conodonts, are generally considered to have special utility for making stratigraphic alignments, locally and globally, but because of relative ease of dispersal are customarily assumed to have minimal value for biogeographic analysis. There are, however, instances where restricted distribution of pelagic taxa-in the present instance conodont taxa-may

173

accord with fonner closer juxtaposition of specific continental margins or provide supporting evidence for global pattems of oceanic circulation. Because of absence from manifestly polar or nearpolar regions, such as the Malvinokaffric Realm of southern Gondwana, conodonts are infelTed to have lived in tropical and temperate waters (Telford, 1979; Klapper & Johnson, 1980).

Although most Devonian conodont genera occur in Australia, some-when they do OCCUf

are relatively rare numerically or occur atvety fe,,, localities. At species-level there are salient occurrences of Early Devonian forms with limited geographic distribution (Table I). This interval has been long recognized as having had provinciality increase to a maximum in early Emsian times and thereafter decline (e.g., Boucot et ai. 1967, 1969; Talent, 1972; Klapper & Johnson, 1980; Boucot, 1975; Oliver & Peddel; 1979, 1989). Contrary to this pattem, Flessa & Hardy (1988) on the basis of cluster analysis of Devonian conodonts, concluded that provinciality during the early Lochkovian was high but declined during the late Lochkovian and Pragian, then increased to a maximum in the Emsian. It should be noted, however, that only 5 localities were used for the analysis for the pesavis and sulcatus zones. The database (Figs 5, 6; Table I), being now much more extensive, calls for reconsideration.

Many conodont genera are either not present or are found rarely in Australia. The genus Sannemannia AI-Rawi, for example, has not been reported from Australia, and Pandoricriodus Mawson, Talent & Furey-Greig is as yet known only from E Australia (Mawson, ef ai., 1995; Druce, 1970b). Biofacies considerations notwithstanding, Icriodus Branson & Mehl does not occur in large numbers prior to the late Emsian (Mawson, 1986; Mawson et ai., 1988, Mawson & Talent, 1989); species of Pelekysgnathus Thomas occur even less frequently-from the late Famennian of the Canning and Bonaparte Gulf basins, Westem Australia (Glenister & Klapper, 1966; Druce, 1969; Seddon, 1970; Metzger, 1994), with only one confirmed occunence from EAustralia: from NE Queensland (Telford, 1975). Representatives of Neopanderodus Zeigler & Lindstrom are known from Australia, the W Camic Alps of Austria (Schonlaub & Flajs, 1993), the Armorican Massif, France (Lardeux & Weyant, 1993), the Rhenish Slate Mountains, Germany (Ziegler & Lindstrom, 1971), W Turkey (Ziegler & Lindstrom, 1971), the Urals (Khodalevich & Tschemich, 1973) and Alaska (Huddle in Churkin

Fig. 5. Coverage of conodont data for the Early and Middle Devonian ofE Australia (from Mawson & Talent, 2000). For correlations between key Early and Middle Devonian sequences see Appendix 2.

174 AAP Memoir 23 (2000)

Q

M/\ >


& Brabb, 1968). Apati from the Rhenish Slate Mountains and the Urals, all a", from thenOlihem margin of Gondwana. The early Late Devonian Playfordia Glenister & Klapper appears to be restricted to the Canning Basin, WestemAustralia (Glenister & Klapper, 1966; Druce, 1976), the Rhenish Slate Mountains, Germany (Bischoff & Ziegler, 1957) and Alberta, Canada (Uyeno, 1974).

The present distribution of a number of conodonts may be taken as indicative of fonner close juxtaposition of the crustal blocks, or of circulation pattems facilitating migration between these blocks. Kimognathus Mashkova, a lllonospecific conodont genus, for example, is recovered typically from the pesavis Zone in terranes now located in Australia, Tajikistan, Uzbekistan and Arctic NOIih America. Such ties are consistent with restriction of the very earliest polygnathids - Polygnathus trilinearis (Cooper) and P. zeravshanicus (Bardashev & Ziegler) - to crustal blocks in EAustralia and Tajikistan. These ties persisted through the kindlei and pireneae Zones of the Pragian. By the pireneae Zone, polygnathids, specifically the zonal species P. pireneae Boersma, had developed a wider distribution extending from E Australia (Mawson et al., 1992; Mawson & Talent, 1994a; Talent & Mawson, 1999) and Tajikistan (Bardashev, 1991; Bardashev & Ziegler, 1992) through other regions on or close to the n011hern margin of Gondwana - Spain (Boersma, 1974; Valenzuela-Rios, 1994) and South China (Hou et al., 1988)-as well as in central Nevada (Murphy & Matti, 1983), the Klamath Mountains, western North America (Savage, 1977) and Baltica, specifically in the Frankenwald, Germany (AI-Rawi, 1977). Restriction of P. hindei Mashkova & Apekina to crustal blocks in Australia (Wall et al., 1995),

175

Tajikistan (Bardashev, 1991; Bardashev & Ziegler, 1992) and Uzbekistan (Mashkova & Apekina, 1980; Yolkin el al., 1989) accords with persistence of old linkages into the early Emsian. During the early Emsian, Ozarkodina prolata Mawson occuned in large numbers both in Australia (e.g., Mawson, 1987a) and at La Grange, northwest France (Bultynck, 1989), another region believed to have formed part of the nOlihem margin of Gondwana during mid-Palaeozoic times.

From late in the Emsian there was a proliferation of polygnathids (Klapper & Johnson, 1980). During the early Middle Devonian, endemism in polygnathid-dominated faunas reached its acme with 68% endemism in the costatus Zone and 74% endemism in the GUsh'aUs Zone (Klapper & Johnson, 1980). Thereafter endemism diminished with only a few species such as Polygllathus parawebbi having a somewhat restricted distribution (Australia, Nevada, eastcentral Alaska, SE Alaska, Northwest Territories and the Russian Platfonn). From varCllS Zone onwards, there was a marked decline in endemism. By the Frasnian, endemism was reduced (e.g., Flessa & Hardy, 1988) but conodont faunas were by no means cosmopolitan. Klapper (1995), using the Raup & Crick (1979) Probability Index of Similarity with well-documented conodont faunas from the Frasnian of the Canning Basin, the Montagne Noire, United States, Canada and the Russian Platfoml, demonstrated that at least a third of known Frasnian conodont species were endemics.

Famennian conodont faunas were highly cosmopolitan, but quantitative data in suppOli of this conclusion are lacking. Palmatolepid faunas inhabiting continental slope and basinal environments appear to have been ubiquitous. Australian endemics at this time include Icriodus

Fig. 6. A selection of Devonian conodont species believed to be provincially restricted (cf. Table 1): A, Ozarkodbw pseudomiae Mawson & Talent. Lateral view of NMVP 142127, SL28, The Basin, E Victoria, x60. B, C, Pandoricriodus imparilis Mawson, Talent & Furey-Greig, B, lateral view ofUQY 7694, Duncan 9, Horrigan Creek, Queensland, x 55. C, upper view ofUQY 7693, same locality as preceding illustration, x 60. D, Ozarkodina linearis (Philip). Lateral view ofNMVP 99037, BON60.5-65, Bonanza Gully, Bindi, Victoria, x 45. E, Ozarkodina prolata Mawson, Lateral view of AMP 117131, SR245, Sawpit Ridge, Bindi, Victoria, x 45. F, Bipennatus palethorpei (Telford). Upper view ofUQY 4023, JES6--7, Jesseys Springs, Broken River, NE Queensland. G, Kimogllathus alexeii Mashkova. Upper view of AMF 104971, 'Canobla', Stuart Town, NSW, x75. H, P. zeravshanicus (Bardashev & Ziegler). Upper view ofAMF 104937, NlTh1113, 'Canobla', Stuart Town, NS\V, x 60. I, Polygnathus trilinearis (Cooper). Upper view ofN1'vfVP 142100, BF4, Boulder Flat, E Victoria, x 60. J, Polygnathus pireneae Boersma, Upper view of AMF 104914, NIN122, Red Hill, NS\V, x 60. K, P. zeravshaniclis (Bardashev & Ziegler). Upper view of AMF 104910, NIN3, Red Hill, NSW, x 75. L, Polygnathus gilklapperi Mawson & Talent. Upper view of AMP 90192, OKE2.1-2.2,Attunga, NSW, x 75. M, Polygnathlls pugiullculus Mawson. Upper view ofUQY 3983,JES9, Jessey Springs, Broken River, Queensland, x60. N,Icriodus sclariJomzis Mawson & Talent. Upper view ofUQY7814, 'Dotswood " NE Queensland, x90. 0, P. "'I'yattiMawson & Talent. Upper view ofUQY7787, W1vfP 504, Mount Podge, Queensland, x45. P.Polygnathus hieroglyphica Mawson & Talent. Upper view of UQY 7772, \VMP 504, Mount Podge, Queensland, x30. Q. Polygnathus costulifera Mawson & Talent. Upper view ofUQY 7776, EMF 48, Mount Podge, Queensland, x60.

176 AAP Memoir 23 (2000)

GENUS LOCALITY REFERENCE

Australia Mawson et aI., 1988; Wilson, 1989; Sorentino, 1989; Brock, 1995; Druce, 1971

Kimognathus SE Alaska Savage & Gehrels, 1984

alexeii Zeravshan Range Mashkova, 1968; Bardashev, 1991

lien Shan Bardashev, 1991; Bardashev & Ziegler, 1992

Urals Sapelnikov et aI., 1995

Australia Cooper, 1973; Mawson et aI., 1992; Mawson & Talent, Polygnathus 1994a; Talent & Mawson, 1999 trilinearis

Shishkat, Tajikistan Bardashev, 1991; Bardashev & Ziegler, 1992

Polygnathus Australia Mawson et aI., 1992; Talent & Mawson, 1999

zeravshanicus Shishkat, Tajikistan Bardashev, 1991; Bardashev & Ziegler, 1992

Australia Mawson et aI., 1992; Mawson & Talent, 1994a; Talent & Mawson, 1999

Pyrenees, Spain Boersma, 1974; Valenzuela-Rios, 1994

East-central Alaska Lane & Ormiston, 1979; Savage et al., 1985

Bathurst Island Uyeno in McGregor & Uyeno, 1972

Polygnathus 'rukon Lane (in Ziegler (ed.), 1977; Savage et aI., 1985

pireneae Frankenwald, Germany AI-Rawi, 1977

Klamath Mountains Savage, 1977

Central Nevada Murphy & Matti, 1993

South China Hou et aI., 1988

Zinzilban, Zerafshan Yolkin et aI., 1989


Australia Wall et al., 1995 Polygathus Zinzilban, Uzbekistan Mashkova & Apekina, 1980; Yolkin et aI., 1989 hindei

Shishkat, Tadjikistan Bardashev, 1991; Bardashev & Ziegler, 1992

Australia Mawson & Talent, 1994b Polygnathus Nevada Klapper, 1977 gilk/apperi

East-central Alaska Lane & Ormiston, 1979

Australia Philip, 1966; Flood, 1969; Pedderet al.,1970; Mawson, 1987 a

Ozarkodina 'rukon and Ellesmere Fahraeus, 1971 linearfs Island, Canada


Ozarkodina Australia Mawson, 1987a

pro/ata La Grange, NW France Bultynck, 1989

Bipennatus Australia Telford, 1975; Pickett, 1978; Mawson et aI., 1985; palethorpei Mawson, 1987b; Mawson & Talent, 1989

Table 1. Endemicity of selected Early Devonian (Pragian/Emsian) conodont species with areas of occurrence and relevant reference.

sclarifonuis Mawson & Talent, Polygnathus wyatfi Mawson & Talent, P. costulifera Mawson & Talent and P. hierogl)phica Mawson & Talent (Fig. 6), whereas P. karadjalis Vortonsova & Kuz'min has been found only in E Australia and

central Kazakhstan (Mawson & Talent, 1997). Conodont distribution thus indicates, in general,

that the Early Devonian was a time of marked provincialism, reaching a climax in the costatus and australis zones ofthe Eife1ian. It is suggested


that the distribution of conodonts in the Pragian suggests either closer juxtaposition of Australia with the Khodzha-Kurgan area (ZeravshanRange) of SE Uzbekistan and SE Alaska, or provides suppOlting evidence for global pattenlS of oceanic circulation that might facilitate ease of transpOlt between these areas. During latest Pragianearliest Emsian, connections with Spain and South China appear to have been made. Through the late Eifelian to the mid Givetian ( varcus Zone) there was a waning of provincialism suggesting closer proximity of crustal blocks or a mOTe effective pattern of ocean currents transpOlting larvae or planktonic taxa. Coinciding with the onset of a major worldwide regression cycle (Johnson et al., 1985) in the varcus Zone, conodont faunas took on a morc cosmopolitan aspect although faunas retained up to 30% endemism in the Givetian, a period when numerous transgressive events took place (Johnson et al., 1985). Waning of pfovinciaHsm from the Frasnian to relative cosmopolitanism in the Famennian suggests that a numberofcmstal blocks were now in very close contact and/or the ocean currents were such that pelagic organisms were easily conveyed from place to place. However the degree of similarity of faunas of the Famennian remains to be tested quantitatively for non-palmatolepid faunas.

Ammonoidea (R.T. Becker) The spatial distribution of Devonian

ammonoids is still rather poorly known although palaeobiogeographic differences of pelagic groups give distinctive signals for routes of faunal exchange between open shelf areas of major clUstal blocks. Both very shallow marine seaways and deep oceanic areas may have acted as balTiers for ammonoids characterised by constraints for maximum vertical migration in the water column and by sensitivity for high water agitation caused by storms, etc. Although many taxa were pan( sub )tropical, the impOltance offacies control on Devonian ammonoid distribution has become lncre apparent in recent years. Trends towards endemic evolution in open seas appear to be a clearer indication of plate tectonic isolation of regions than endemism in small niches of benthic and nearshore groups. Improved time resolution of ammonoid occurrences gives prospects for a better-refined history of palaeobiogeographic changes.

Global diversity analyses of all Late Devonian ammonoids at the generic level, based on the best available time resolution, have been undertaken by Becker (1993). This study gave clear indications of relationships between eustatic changes and times of diversification and extinctions ("species area effect"). So far, there

177

have been no published investigations either at species level or on a regional scale, directed towards elucidating differences between lower and major taxonomic levels, or to establish differences betweenregional and global diversity patterns. Of special interest are links between diversity changes and the degree of endemism; both were influenced by sea-level changes.

Special interest attaches to the earliest goniatites from zones low in the Taravale Formation of the Buchan area, E Victoria: Teicherticeras desideratum (Teichelt) fi·om the earliest Emsian perbonus Zone and Talenticeras ta'eJlti Erben from high in the dehiscens Zone. They OCClli" in association with bactlitids: Bach"ites· hOl-vitti Teichert, Lobobactrites inopinatlls Teichert and other bactritids (Teichelt, 1948). The goniatites, dated by conodonts (Mawson, 1987a), are among the oldest ammonoids in the world. Teic/terticeras is a semi-cosmopolitan genus.

Late Devonian Callning Basin ammonoid palaeobiogeography alld diversity. Principles of Devonian ammonoid palaeobiogeography have been established by House (1964, 1973). Palaeobiogeographic relationships of ammonoids from the Famennian of Western Australia (Canning Basin) were discussed by Petersen (1975). He found strong similarities with other, widely separated, regions that formed part of the Late Devonian Prototethys. Intensive new field work in the area since 1989 has greatly expanded knowledge of ammonoid taxonomy and stratigraphy of the region (Becker et al., 1993, Becker & House, 1997). The Canning Basin has the most diverse ammonoid succession in the world, comprising a total of 175 taxa from the Frasnian (UD I-A/B) to middle part of the Famennian (UD N)----a formidable basis for a reappraisal offaunallinks with other regions as well as for inter-regional diversity analyses.

Canning Basin ammonoid taxa fall in three categories conceming their spatial distribution: ( a) taxa known from several to many widely separated areas (semi-cosmopolitan to cosmopolitan), (b) endemic taxa, and (c) "spot taxa" with patchy distribution in two widely separated regions without evident intermediate occun-ences. The latter are especially interesting for establishment offaunallinks.

There are strong differences at various taxonomic levels. Australia has no endemic Devonian ammonoid family and, at the generic level, endemism is also low (around 5%). Diversity is higher in the Frasnian (35 genera/subgenera with 103 species/subspecies) than in the Famennian I1-IV (32 genera with 72 species). The average diversity of Frasnian zones lies at ca. 9.5 species

178 AAP Memoir 23 (2000)

G H


but only at ca, 7 species for Famennian zones. These figures are based on real occunences and do not include regional "Lazams episodes" (record gaps) oftaxa. Bactritids have been omitted since their record is strongly biased by preservational aspects (easy species identification only in haematitic faunas). Faunas are generally not as diverse as in some Gemmll or Polish submarine seamount sections. Several diversity peaks clearly conelate with transgressive episodes, but not all eustatic rises allowed diversification or the spread of more species into the Canning Basin (see Becker & House, 1997). The maximum diversity with 31 species/subspecies was reachedio the Virgilloceras erraticum Zone (UD I-J2). Other maxima correlate with the peak hypoxic phase ofUD 1-F2, with the "semichatovae Transgression" (UD I-II), and with the Annulata Event (UD IV-A). Regionally, diverse faunas also have been found in the latest Frasnian (UD I-Lla), during the regressive period after the Lower Condroz Event (rhomboidea Zone, upper part ofUD II-D), and at the base of UD III. Some levels such as UD 1-K, UD II-G2, and UD III-A3 are probably still insufficiently sampled and give unusual diversity minima. Whereas the Lower Kellwasser Event had a drastic effect in the Canning Basin (regional extinction of 27 species), there was a gradual decline of ammonoid faunas towards the end of the Frasnian. However, not one species survived the Upper Kellwasser Event, not even tornoceratids as 'Lazarus taxa', and the Pa. triangularis to Middle crepida Zones (UD II-A! B) are devoid of ammonoids (Becker et al., 1991).

There is no distinctive palaeobiogeographic difference between the two Late Devonian stages. The global Kellwasser Crisis at the end of the Frasnian had no lasting effect on faunal exchanges. 'Spot taxa' give strong links with Germany, with the Timan and, surprisingly, with E North America but not with North Africa.

At the species level, endemism is significant and around 50%, both in the Frasnian and in the

179

Famennian. Analysed zone by zone, the average rate of endemism lies at 40.5% in the Frasnian and at 42.3% in the Famennian. Many still undescribed forms, however, are closely related to taxa from Gennany, Poland, E North America, and the Timan. Low diversity faunas have a higher percentage of endemic fonus and seem to contain specialists adapted to 10caVregional environmental conditions unfavourable for most ammonoids. Relatively low rates of endemism « 20% of taxa) were found during some specific but not during all transgressive episodes. Examples are the basal Rhinestreet eustatic pulse (UD I-GI), the 'semichatovae Transgression' (UD I-H), and the two eustatic rises during the marginijera Zone (UD II-F2 and II-GI). It is suggested that differences in ammonoid diversity and endemism during individual transgressive phases reflect Vatying overprints by basin subsidence histmy and by associated regional ecological developments.

There are no strong faunal similarities between NW Australia and South China but the latter region has so far yielded only rather sparse faunas from the same Late Devonian time intervals. In the Famennian II and IV, the Canning Basin and NSW have no species in common. Both regions may represent different palaeobiogeographic realms. EastemAustralianammonoid source areas are part of tenanes, which may have attached later to the craton, or there may have been some balTier to dispersal of faunas from W to E or vice versa. There was certainly also a facies influence on different faunal composition since Canning Basin UD IV -assemblages (piker Hills) were found in a rather unusual hemipelagic shallow black shale facies. NSW faunas (Jenkins, 1966, 1968; Wright et al., 1990) contain both endemic species (e.g., 'GeJluclymenia keepitensis') and fonns known from W Europe and NOlih Africa.

Significant Frasnian 'spot taxa' ofthe Canning Basin areProchorites alveolatus andProbeloceras ltilheri (link to New York State), the Costamanticoceras koeneni and <MateJ'Jlocel'as J

Fig. 7. A selection of Australian Devonian chitinozoans believed from available data to display a high level of provinciality: A, Bulbochitina bulbosa Paris, UTGD 127068, Point Hibbs Fonnation, Tasmania, X350, Pragian, Mndlei Conodont Zone. B, Eisenackitina subdUiva \Vinchester-Seeto, AMF117128, Taravale FOlmation, northeastem Victoria, X350, Emsian,perbolllfs Conodont Zone. C, Calpichitina velata (Wrona),AMF117129, Gana Limestone, central NS\V, X400, Lochkovian-Pragian, pesavis-sulcatus conodont zones. D, Bursachitina (Amplichitina) bidawal Winchester-Seeto, NMVP138573, Taravale Formation, northeastern Victoria, X400, Emsian, perbollus Conodont Zone. E, Gotlandochitina milallensis (Collinson & Scott), M1F117130, Gogo Fonnation, Canning Basin, X350, Late Givetian. F, Bursachitina (Bursachitina) kllrritgo Winchester-Seeto, NMVP138581, Taravale Formation, northeastern Victoria, X300, Elllsian,perbonus-serotinus conodont zones. G, Angochitina comosa Taugourdeau & Jekhowsky, AMF82458, Garra Limestone, central NS\V, X400, Pragian, sulcatus Conodont Zone. H, Angochitina hypenetes \Vinchester-Seeto, AMF8246 I , Garra Limestone, central NSW, X250, Lochkovian-Pragian,pesavis-sulcatus conodont zones. I, Gotlandochitina viridarillm \VinchesterSeeto & Paris, GSWAF49401, SadlerFonnation, Canning Basin, X350, Early asymmetricus Conodont Zone.

180

Late Lochkovian

Pragian

Emsian


SPECIES GENUS

Fig. 8. Number of taxa of Chitinozoa common to Australia and otherregiolls listed for species and genus fcreach stage of the Early Devonian (from \Vinchester-Seeto & Paris, 1995). Number in parentheses refers to the number of species or genera present at each location. Numbers inside the circles are the number of common taxa.

prumiense Groups (link to Rhenish Massif), Manticoceras evolutllm (link to NOIihAfrica), and rare M latisellatllm and M solnzevi (link to Timan). Famennian examples are Torula n. sp., some cheiloceratids (link to Poland), Protactoc/ymenia krasnopolsld (link to the Urals), Karac/ymenia n. sp. juv. (link to Polar Russia), and MaelIeCeraS miller! (Famennian link to New York).

Generally, Wand E Australian Devonian ammonoid faunas can be construed as representing discrete branches of Prototethys with fast faunal exchanges between distant regions. Migrations took place on a W-E route between NW Australia,

the Urals-Timan seaway, and, via PolandGenuany (the Rhenohercynian seaway) to the Appalachians. The high level of species endemism reflects the vast distance between EuropeaniNorth American regions and Australia. Increased knowledge of poorly studied faunas from the Middle East (e.g. Iran) is clUcial for understanding of migration patterns along the northern Gondwana margin mid-way between the wellstudied regions.

Chitinozoans (T. Winchester-Seeto) The database for Australian Early Devonian

chitinozoans is relatively sparse. Inferences based


on it should therefore be viewed as tentative, with some possibility that similarities between Australia and the rest ofthe world may be underestimated.

In contrast to patterns of distribution in the Ordovician (Ordovician chapter herein), Early Devonian chitinozoans display only a moderate degree of provincialism (Figs 7, 8). Recent biostratigraphic studies of Early Devonian chitinozoans from Australia (Winchester-Seeto, 1993a, 1993b) have demonstrated the potential for intercontinental correlation, based on the frequent presence of index and other key species - identified in biozonations based on SW Europe (Paris, 1981). Nonetheless, statistical analysis of whole assemblages has shown that there is a low level of similarity at the species-level between Australia and the rest of the world for late Lochkovian and Pragian faunas (WinchesterSeeto, 1997). Endemicity reached a peak in the Emsian (perbOll11S to sera tin liS Zones) with diverse, but apparently unique assemblages from the Taravale Formation of the Buchan area ofE Victoria (Winchestel'Seeto, 1996). Reasons for the remarkably high level of endemicity are not known. There arc, however, only five published studies on chitinozoans for this time-interval globally. Some ofthe seemingly endemic species from the Taravale FOlmation may also occur in Canada (E. Asselin, pel's. comm., 1997). The actual level of endemicity may therefore be somewhat lower than might presently appear. Six (32%) ofthe chitinozoan genera presently known to have existed in the Early Devonian have not been found in Australian strata: Arrnoricochitina, Cing/uochitina, Linochitina, Margachitina, Urnoc!titina and Urochitina; phylogenetic trees proposed by Paris (pers. comm., 1995), incidentalIy suggest close relationships between five of these genera.

Spot samples from the Middle and Late Devonian shows salient similarity between: I. the ?Iate Givetian portion of the Sadler and Gogo fonnations of the Canning Basin, and the Cedar ValIey Limestone ofIowa and the Hamilton Formation of Ontario; 2. the early Frasnian segment of the Sadler and Gogo formations of the Canning Basin, and the lower La Serre FOlIDation, Montagne Noire, S France (Winchester-Seeto & Paris, 1995); 3. the late Frasnian Mostyn Vale Formation ofthe Tamworth Belt of NE NSW and the lower La Serre Fonnation, Montagne Noire, S France (WinchesterSeeto & Paris, 1995).

There is no obvious explanation for endemism of particular species or genera of chitinozoans. Distributions of genera and/or species do not conform to patterns possibly reflecting palaeolatitudes. Distributions, when placed on a

181

A B c Fig. 9. Key Early Devonian graptolites from SE Australia: A, Monograptus thomas; alexandraensis Jaeger, x2.S. B, Ai thomasi thomasi Jaeger, xl.S. e, M aequabilis notoaequabilis Jaeger & Stein, x2.5. (all from Jaeger, 1988).

plate reconstruction (Scotese & McKerrow, 1990), could have been cOllllected with palaeooceanic currents-in much the same way as the distribution of modem zooplankton is connected with the pattern of contemporary oceanic CUlTents.

Gl'aptolithina (R.B. Rickards & A. I. Wright) The database for Australian Early Devonian

graptolites-recently reviewed by Rickards & Wright (2000}-although small and coming from a vety few scattered horizons, is slowly expanding2.

There are no exclusively endemic forms (Fig. 9), so little can be said regarding the biogeographic linkages ofthe few fonns desclibed to date (Jaeger, 1988; Rickards & Wright, 2000). Jaeger (1966), for instance, described Monograptus thomasi thomasi from several Pragian localities in central Victoria; it is also known from SE Asia and S Tibet-regions that were formerly part of the Gondwana margin-but is also known from Kazakhstan, Arizona, British Columbia and elsewhere (Jaeger, 1966; cf. Carey & Bolger, 1995; Jaeger & Stein, 1969; MUlphy & Beny, 1983; Mu & Ni, 1975; Jaeger, 1983, 1988). Similarly,

182

M aeqllabilis noloeaqllabilis Jaeger & Stein, widespread in east-central Victoria, has proved to be widely distributed outside Australia: inler alia, in Alaska, Bohemia and Bunna.

Tentaculitida (Dacryoconarida) (J.A. Talent) That daClyoconarids have considerable value

for establishing inter-regional and even intercontinental correlations is well known. A major exercise in taxonomy of the pelagic tentaculitids (dacryoconarids), including many faunas Ii-om E Australia, has been undertaken by G.K.B. Albelti over many years, especially over the past 15 years (Albelti, 1987, 1988, 1993, 1995, 1997a, 1997b, 1998, 1999, 2000). Forms identified from the latest Lochkovian to early Emsian from many areas in central and E Victoria and from latest Emsian (sel'Otinlis Zone) to Eifelian of the Broken River region, NE Queensland, have proved to be specifically identical or closely comparable with forms from Morocco, Bohemia and the Harz Mountains of Gemlany. This is not unexpected as all these regions were fOlmerly part of the northern Gondwana continental margin. Elaboration of the bed-by-bed stratigraphic distribution ofthe Australian daclyoconarid taxa and their biogeographic inlplications is awaited.

Brachiopoda (lA. Talent) Early Devonian (Lochkovian--early Emsian)

brachiopod faunas ofE Australia are abundant and diverse, especially in SE Australia. Australian Middle Devonian brachiopod faunas are very poorly known, apmt from small Givetian faunas, principally with stringocephalids, ii-om a small area nearthe N margin ofthe Canning Basin of West em Australia, from the Moore Creek and Burdekin Limestones of NE NSW and NE Queensland respectively, and from the Papilio Formation of the Broken River region of NE Queensland.


Diverse Late Devonian (middle and latest Famennian) brachiopod faunas occur in the BurdekinBasin ofNE Queensland but, apaIt from the, productids, are still undocumented. Late Devonian (plincipally Frasnian) faunas have been documented from the three sedimentmy basins in Westem Australia having Devonian sequences; all brachiopod faunas documented from these three regions have been assumed to be relatively cosmopolitan at generic level. Less is known regarding the Late Devonian brachiopod faunas of eastern Australia, especially the low diversity faunas from the VCly shallow clastics (Lambie Province; Fig. 3) ofSE Australia.

Early Devonian. The Tasman Region ofBoucot el al. (1967, 1969; Boucot, 1975; cf. Fig. 10), proposed for Em'ly Devonian-Eifelian brachiopod faunas from E Australia and New Zealand, is derived from !he Late Silurian and ?Wenlock SinoAustralian Province of Rong el 01. (1995). A distinct New Zealand Province is apparent for Early Devonian brachiopod faunas especially the 'Reefton' fauna. Endemics in the latter include the chonetidine Allaneles Boucot & Johnson, the rhynchonellidine Tallerhynchia Allan, and elements otherwise characteristic of the Malvinokaffric Realm, specifically the terebratulidinePleul'Olhy",l/a Boucot el al. (Allan, 1935, 1947; Boucot el al., 1963; Boucot & Johnson, 1967). The leptocoeliid Auslralocoelia BOllcot & Gill, also characteristic of the Malvinokaffric Realm, is present in the Bell Shale ofNE Tasmania asA. polyspera (Gill) (Boucot & Gill, 1956); it is anticipated to have been present in the New Zealand Early Devonian faunas' (see below). The strophomenid Maol'isll'ophia Allan, Oliginally described ii-om the ?Pragian at Reefton, New Zealand as a stropheodontid (Allan, 1947), is a common form in shallow water Lochkovian-

Fig. 10. A selection of species representative of Early Devonian (early Emsiall) brachiopod genera from Taemas, NS\V (from Chatterton, 1973), believed from available data to be endemic to E Australia or restricted to few provinces: A-D, Malu1'Ostrophia jlabelficauda Campbell & Talent. A, brachial valve interior view of ANU l8933e, x2. B, pedicle valve interior view of ANU 18933d, x2.4. C. enlarged portion of pedicle valve interior of ANU 18933a, x5. D, anterior view of pedicle valve ANU 18925, x2.3. E-J, Chattertonia campbelli (Chattet10n). E, extemal view of pedicle valve epc 10569, x1.8. F, Lateral view ofANU 18951e, x2.5. G, interior view of pedicle valve ANU 18951k, x4. H, interior view of pedicle valve ANU 18951 kx4. T, interior view of pedicle valve ANU 18951d, x4. J, lateral viewofANU 18950, xl.8. K-N,Adrenia e.rpansaChatterton. K, enlarged illustration of brachidium. L, Ventrolateral view of ANU 18985d showing partial preservation of brachidium, x 4.6. M, lateral view of ANU18986, x3.6. N, exterior view of brachial valve cpe 10628, x5.4. O-Q, Cydimia robertsi Chatterton. 0, Anterior view of ANU 18992m x3.6. P, exterior view of brachial valve AND 18990, x5. Q, enlarged illustration of brachidium. R-U, Howittia howitti (Chapman). R, interior of portion of brachial valveANU 18962h, x5.25. S, exterior view ofpedic1e valveANU 18962amxlA. T, anteriolateral view of pedicle valveANU I 8962c, x2. U, interior view of brachial valve ANU 18962d xl.? V-X, Spinella yassensis (de Koninck). V, enlargement of anterior portion of pedicle valve showing micro-omament, x5. \V, posterior and dorsal views of ANU l8966d, x2. X, dorsolateral view of pedicle valve of ANU 18966e, x2.

AAP Memoir 23 (2000) 183

A c

B

F G

I . J R

M N o p Q

s v T X

w

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Pragian and very rarely earliest Emsian clastic sequences in SE Australia, including Tasmania (Gill, 1952). Highly characteristic of the Tasman province is the abundance and diversity of notanopliids, especially in mudrocks and occasionally in limestones (Gill, 1969; Lenz & Johnson, 1985b), though the family was widely distributed elsewhere in the Old World Realm during Early Devonian times fi'Om western Europe (Jahnke, 1971), to the South China and MongoloOkhotsk Provinces and the terranes in WYunnan (cf. Gratsianova & Shishkina, 1977; Su, 1980; Talent et al., 1985,2000; Jahnke & Shi. 1989).

Lochkovian. The very shallow water faunas of the thick clastic late Ludlow-?early Lochkovian McIvor Sandstone' of the Heathcote-Redcastle area' of central Victoria (Talent, 1964) are dominated by rhynchonellidines and have a strong 'cosmopolitan' aspect- atrypidines, especially Allstralina, dalmanellidines resembling Salopina, small spiriferidines resembling HOlVellella and leptaenids-but persistence of the retziaceans Molongia and entry of Maoristrophia herald the increasing provinciality of SE Australian brachiopod faunas through the latest Silurian into the Lochkovian and Pragian of the Tasman Province.

Unit I of the Mount Ida Formation (the Comella Member) is characterised (Talent, 1964) by an assemblage of generically cosmopolitan forms: dahnanellidines including Salopina, and Isort"is (Tyersella), and the stropheodontid Leptostrophiella; the retziaceans Molongia and Athyrisina appear to be restricted to the Tasman and South China biogeographic provinces (cf. Rong et al., 1995).

Unit 2 of the Mount Ida Formation (the Dealba Member, formerly the Notoconchidium Beds)--characterised by often great abundance of the cuboidal rhynchonellidine N. tasmaniense (Etheridge), originally describedasN. thomasiby Gill (1951a)---has generally low diversity faunas with cosmopolitan genera including the orthidine Skenidioides (Boucot et al., 1966), the sowerbyellid Plectodonta, and nondescript species of the small spiriferidine HOlVellella, poorly preserved atrypidines, and, rarely, the strophomenidMaoristrophia.

Unit 3 of the Mount Ida Formation, the Stoddart Member (fOlmerly the PleurodictYllm Beds; cf. Edwards et al., 1998), has a diverse fauna of somewhat larger brachiopods (Talent, 1964), notably the leptaenids Notoleptaella tingu(fera Gill and N. otophera Gill, the strophomenid Maoristrophia banksi Gill, the stropheodontids Mesodollvillina (Mesodouvillina) limbimllra (Talent) and Strophonella manta Talent, and the notanopliid BOl/cotia australis (Gill). Associates


are presently unidentifiable species of the dalmanellidines Isorthis (Tyersella), Rhipidomella? and Salopina?, rare plectambonitids (Plectodollta), occasional leptaenids including Leptagonia?, relatively connnon stropheodontids including Leptostrophiella, various rhynchonellidines, atrypidines including Alistratina, athyrididines including Meristella and Nucleospira, the retziidine Molongia, and the spiriferidines Howellella and Havlicekia.

The brachiopod-dominated faunas of the McIvor Sandstone and Mt Ida Formation have much in common with the ?Pridoli and Lochkovian of the Winduck and Amphitheatre groups of The Meadows area, southwest of Co bar and the Bogan Gate-Tmndle area of western NSW (Sherwin, 1990,1995; Fiildvaty, 2000). Sherwin (Ioc. cit.) has identified Retziella capricornae (McKellar) and species of Salopina, Isorthis (Protocortezorthis), Mesodollvillina (Mesodollvillina), Iridistrophia, Sphaerirhynchia, Aliypa, Howellella, apparently Howellella (Hysterohowellella) jacqueti (Dun) from the Meadows area. Similar faunas have been described from the Cookeys Plains and ConnemalTa Fonnations ofthe Bogan Gate-Trundle area of central NSW. Preservation of these faunas is poor but, with the exception of the S. pittmani fauna, they are strongly reminiscent of the faunas of the Mt Ida FOlmation of central Victoria. A younger, less diverse fauna with Spinella pittmani (Dun) occurs in the Troffs FOlmation and Jemla Limestone Member of the Gleninga Formation of the same region; it is Pragian but may extend into earliest Emsian (Fiildvary, 2000).

What may be construed as a representing a marine or quasi-marine community representing a shallower enviromnent than preserved in the Mount Ida FOlmation occurs in the Silverband Fonnation within the predominantly non-marine Grampians Group of W Victoria (Talent & Spencer-Jones, 1963). The brachiopod fauna is monospecific consisting of the nondescript inarticulate brachiopod Lingula borungensis (Chapman) occulTing sometimes as pavements, presumably generated by intertidal or velY shallow subtidal sluicing. Associated with it are low diversity thelodont-acanthodian-ostracod faunas (Talent & Spencer-Jones, 1963; Turner, 1986). The horizons are thought fi'om radiometric data to be about Lochkovian (Early Devonian) or perhaps somewhat younger in age (Mawson & Talent, 2000). The low diversity fauna does not appear to be biogeographically significant.

The 'Yeringian' brachiopod fauna of the Humevale Siltstone, long assumed to be Silurian but first demonstrated to be substantially Early Devonian by Gill (1942), is more diverse than


brachiopod faunas of the Mount Ida Formation (patticularly Unit 3, the Stoddart Member)'. The most diverse Humevale brachiopod faunas are fi'omLochkovian horizons (late Lochkovian, and conceivably even earliest Pragian) in the Mooroolbark-Lilydale-Coldstream area E of Melbourne, notably at Mooroolbark and Chimside Park, N of Lily dale. Most of the upper Humevale brachiopod munas have yet to be described, but typical elements are the long-ranging plectambonitidPiectodonta bipartita (Chapman), the leptaenid Notoleptaena otophera Gill, the strophomenid Maoristrophia keblei Gill, the rhynchonellidine SphaerirhYllchia globularis Talent, the atrypidine Spinatrypa fimbriata (Chapman), the notanopliids BOllcotia australis (Gill), B. withersi (Gill), B. loyolensis Gill and Notanoplia philipi Garratt, the large spiriferidine Plicocyrtina cooperi (Gill) and undescribed or poorly known species of 'cosmopolitan' genera: Schizophoria. Fascicostella, Leptagonia, Leptostrophiella, CostistropllOnella?, Nlicleospira, Cyrtinaella, and various chonetidines, rhynchonellidines, atrypidines and spiriferidines, some of which were named almost a century ago but are in need of revision on the basis of better material than was available when they were described. Brachiopod faunas from about the middle of the Humevale Siltstone have many elements in common with brachiopod faunas fi'om low in the Maradana Shale-about eurekae"sis Zone of the Lochkovian-in the vicinity of Manildra, west-centralNSW (Savage, 1974).

The faunas from the Maradana Shale include Plectodonta bipartita (Chapman), Maoristrophia keblei Gill, Leptostrophiella affinalata (Gill). Rugoleptaena lIndulifera (Talent), Notanoplia pherista Gill, Plicocyrtina cooperi (Gill), forms present in the faunas of the Humevale Siltstone of central Victoria, as well as Spirigerina (Spirigerina) suprommginalis (Khalfm), Isorthis (protocortezorthis) inostranzewi (Peetz) or a fmID very close to it, Molongella talenli Savage, Resserella strllszi, and species of Skenidioides, Dicoelosia, Murijerella, Noto/eptaena, Strophochonetes, Machaeraria, Al1ypina, Al1ypa, Oglu, Australilla, Coelospira, Cyrtin(l, Havlicekia and Quadrithyris.

Lochkovian (ellrekaensis Zone and possibly earliest delta Zone) silicified brachiopod faunas have been described from Windellama (Mawson & Talent, 1999). The faunas consist of: Pelecymya caperata Mawson & Talent, Schizo phoria el'ugata Mawson & Talent, Gypidula pelagica lunata Mawson & Talent, Morinorhynchus fly peter Mawson & Talent, Asynnnetrochonetes picketti Mawson & Talent, Machaerariaformosa (Hall), Hadl'orhynchia? attinarum Mawson & Talent,

185

Sphael'irhynchia? mastodon Mawson & Talent, Atrypa nieczlawiellsisis Kozlowski, Cyrtina praecedens Kozlowski, Ambothyris? inopis Mawson & Talent, Howellella placeotextilis Mawson & Talent, H. alatextilis Mawson & Talent, H. legirupa Mawson & Talent, Reticulariopsis saginatus Mawson & Talent and unidentified forms ofthe dalmanellidine Isorthis and the acrotretids Opsiconidion and Schizotreta. Of 14 species described, all but three were described as new, but all genera, including a new genus Pelecymya, are widely distributed Old World Realm or cosmopolitan genera. Silicified brachiopod faunas associated with abundant Cladopol'a and eurekaensis Zone conodonts (Colquhoun, 1995; Mawson & Talent, unpub. data) occur in the Uppe1TI10st 5 m ofthe Clandulla Limestone in the Kandos area, SE of Mud gee.

Lochkovian (delta Zone) silicified brachiopod faunas have been described from two Garra Limestone sequences in east-central NSW: at Manildra and The Gap (Fmell, 1992). The fIrst ofthese(Savage, 1968a, 1968b, 1969, 1970, 1971; Trotter, 1990; cf. Talent et al., 2000) is from a sequence interpreted as a tongue of Garra Limestone extending westwards into the Cowra Trough. The following fonns have been identified: Dolerorthis packhami Savage, D. smitM Peny, Planicardinia cal'roli Savage, Skenidioides robertsensis Johnson et al., Talentella sub/J1urifer (Johnson et al.), Isorthis (Arcualla) slilcata Wahnsley & Boucot, Eoschuchertella burrenensis (Savage), Anastrophia (Grayina) australis Savage, Allstralirhynchia eudalensis Savage, Eoglossinotoeehia caellminata Havlicek, Aflypino talenti Savage, Ogilviella prolifica Savage, Al1ypa inversa Savage, Reticulatrypa fairhillensis Savage, Spirigerina (Spirigerina) marginaliformis Alekseeva, Meristina? subovata Savage, Cyrtina imbricata Fan'ell, Ambocoelia? dorsiplicata Savage, Rufispil'lfer australis (Savage), Quadl'ithyris molongensis Savage, Prol'eticularia beddei Savage and species of Ptychopleul'ella, Resserella, Isorthis (Protocol'tezorthis), Colletostracia, Iridistrophia, Cymostrophia, Gypiduia, Linguopugnoides, Maehaeraria, Leptathyris, Metista, Nlicleospira aud Cyrtina-all except the recently proposed Co/leiosfl'acia being widely distributed through the Old World Realm. Only six of the 21 species identified have beenrepOlted from regions other than Australia: four from Laurentia, one fi'om Bohemia, one relatively cosmopolitan. A high level of species-endemicity is indicated.

A brachiopod fauna with 16 generically allocated species - 7 of them new - has been reported from an approximately coeval sequence of GaIT a Limestoue atThe Gap, about 30 km NNE

186

of Manildra (Farrell, 1992; cf. Mawson & Talent, 1999; Talent el al., 2000; Brock el al., 1995): Opsiconidioll minor Popov, Dolerorthis angustimusculus Farren, SchizopJlOria? herkioll Farrell, ]ridistrophia mawsonae Farrell, Eoschuchertella blllTenensis Savage, Col/elos/racia roslynaeFalTell, Gypidula pelagica aus/re/ux Fan'ell, Anasll'Ophia (Grayina) australis Savage, Machaeraria catombalellsis Strusz, Ogilviella p/yarion Farrell, Reticulaflypa Jairhillensis Savage, Spirigerina (Spirigerina) supramarginalis (Khalfin), S. (S.) marginaliformis Alekseeva, Ambocoelia? dorsiplicata Savage, Cyrlina praecedens Kozlowski, Howellella talen/i Farrell, and not specifically identified fonns of Opsiconidion, Skenidioides, Mlirijerella, Atl},pina, Atlypa, Megakozlowskiella and Prore/iclilaria. Again there is high level of endemicity at species level, only two ofthem being koown from outside east-central NSW, both being widely distributed fmills of Spirigerina.

From Eurimbla, about 12 km N of The Gap, Brock (1995 ms.) has documented a diverse late Lochkovian pesavis Zone silicified fauna from a thick sequence commencing just above the top of the del/a Zone. Of the 54 species identified, only 5 are known from the rather older sequences farther S at Manildra and The Gap, II are new, and at least 15 occur also in the GaITa Limestone in the Wellington Caves area farther north (see Pragian section below). The fauna is noteworthy for prominence ofthe orthidines Doleror/his and P/ychopleurella, abundance of the rhynchonellidineMachaeraria catombalensis Strusz low in the sequence, and especially for abundance of atrypidines (a dozen species).

The fauna of the Boola Siltstone (Philip, 1962}-[atest Lochkovian and earliest sllica/lls Zone (Mawson & Ta[ent, [994a }-consists ofthe orthidine Doleror/his perscllip/a Philip, the dalmanellidines Par111or/hina impensa (Philip), Isor/his (Tyersella) /ypica Philip and 1. (Pr%cor/ezor/his) Jes/iva Philip, the p[ectambonitidPlectodon/a bipartita (Chapman), the [eptaenid Nololeptaena olophera Gill, the stropheodontacean Sll'Ophonella (Quasisll'Ophonella) gippslandica Philip, the atrypidines Reticula/l},pa Ihomsonensis (Ta[ent), Spirigerina (Spirigerina) supramarginalis sibirica Rzhonsnitskaya and Auslralina lenticulala (Philip), the notanop[iids Soucolia australis (Gill) and No/anoplia philipi GalTatt, as well as species ofthe dahnanellidines Dalejina and Schizophoria, chonetidines, poorly-known pentameridines referred to "Gypidula", and atIypidines including Oglu and the spiriferidine MyriospirijeJ: Like the faunas from the Humeva[e Siltstone and the faunas from the Heathcote-Redcastle and other


areas in central Victoria, this fauna has an abundance of genera that were widespread throughout the Early Devonian in the Old World Realm.

Not well constrained as to age relative to the Silurian-Devonian boundary are various faunas from the Zeehan-Queenstown area ofWTasmania (Gill, 1948, [950) The Florence Quartzite with Noloconchidium lasmaniensis (Etheridge) may be Pridoli; associated fonns are poorly preserved or nondescript (Ta[ent el al., 2000). The overlying Bell Shale (Gill, [950; Boucot & Gill, 1956) appears to be large[y if not entirely Lochkovian by analogy with the "Yeringian" sequences, especially the Humeva[e Siltstone, of central Victoria, but available data are not tightly constraining with respect to the Silurian-Devonian boundaty. The fauna consists ofthe p[ectambonitid Pleclodonla bipartita (Chapman), the strophomenid Maorislrophia keblei Gill, the atrypidines Notanoplia pherisla Gill and Australocoelia po/yspera (Gill)' a species of eospiriferid, Havlicekia, and other fmills for which there is insufficient infonnation.

Two units of the Takaka Terrane of the NW part of the South Is[and of New Zealand are of interest: the Hailes Quartzite below passing gradationally into the Baton Fonnation above. The fOlmer, assumed to be no younger than Pridoli, has produced poorly preselved specimens of the unusual rhynchonellidine N%conchidium (Wright & Garratt, 1991), long koown from the PridoliLochkovian of central Victoria and western Tasmania. The Baton Formation is assumed to be Lochkovian because afthe occurrence oflcl'iodus close to I. woschmidli from below the richly fossiliferous interval (J. Simes, pers. corum., [996). The fauna (Shir[ey, 1938; Ta[ent el al., 2000; for stratigraphic infonnation see Willis, [965, Coleman, [98[ and Bradshaw, [989) is in need of comprehensive reinvestigation. It consists of the dahnanellidine Fascicoslella balonens;s Walmsley & BOllcot,Hipparionyx minor shirleyi Gratsianova & Ta[ent, the spiriferidines Havlicekia secans (BalTande), Plicocyrtina cooperi (Gill) and unidentified or unidentifiable species of Schizophoria, Reeftollia, Maoristl'Ophia, Mesodouvillina (Prolocymoslrophia) and Nucleospira. ill generic composition the fauna is velY similar to Lochkovian-Pragian faunas ofSE Australia, especially the Humeva[e Siltstone and Bell Shale discussed above. What would today be called its Old Wor[dRea[m linkages were earlier noted by Shirley (1938).

Pragian. The Boo[a Siltstones are overlain by sulcalus Zone (but not earliest sulcalus Zone) to early dehiscens Zone [hnestones of the Coopers Creek Formation (Mawson & Ta[ent, 1994a). The


lower limestones ofthis unit ~sulcatl!s to kindlei Zones-have produced a small fauna (Philip, 1962; Brock et al., 1995) with a few taxa cooccurring in the underlying Boola Siltstone: the acrotretids OpsiconidioJl arcticon Ludvigsen, O. minor Popov, O. robustul11 Brock et aI., and Concaviseptulll lauriei, the dalmanel1idines Parmorthina impensa (Philip) and Isorthis (lYersella) typiea Philip, the plectambonitacean Pleetodonta bipartita (Chapman), the notanopliid Notanoplia philipi Garratt and the smooth atrypidine Australina lentieulata (Philip). Also present are species of the strophomenid Maoristrophia, rare large cuboidal rhynchonellidines, and spiriferidines: Myriospirifer and small delthyridids. Noteworthy is the relative abundance and diversity of acrotretids.

At Waratah Bay6 in SE Victoria, impure limestones of the early Pragian (sulcatus Zone) Waratah Limestone, coeval with the Coopers Creek FOlmation limestones, have an undescribed low diversity brachiopod fauna, notably at Robin's Rocks, with a large cuboidal rhynchonellidine resembling Sphaerirhynehia, generally small delthyridids and a rare Olihotetacean. The fauna is strongly Old World in aspect. The unconfOlmably overlying later Pragian (kindlei-pireneae zones) Bell Point Limestone also has a low diversity brachiopod fauna of the athyridines Buchanathyris westoni Talent and Athyris waratahensis (Talent) and the spiriferidine Spinella yassensis (de Koninck) recalling the low diversity faunas ofthe Buchan Caves Limestone ofE Victoria, especially low in that formation.

Brachiopods have been described from limestone olistoliths of probable early Pragian and possibly late Lochkovian age (Mawson & Talent, 1994a) in the Wumtun Formation in Marble Creek and Deep Creek E and SE of Walhalla. These include species ofthe dalmanellidine Dicoelosia, the lepta enid Leptagonia, the stropheodontid Mesodouvillina (Protoeymostraphia)?, the pentameridine Gypi dula, rhynchonellidines including Aseptirhynehia? and Taimynynx?, atrypidines including Reticu!atlypa thomsollensis (Talent) and A/l)pa spp., as well as species of Howellella including H. lirata Talent (Chapman, 1903; Talent, 1956b; Talent et al., 2000). Approximately coeval faunas from clastics at Loyola have produced the notanopliid Boueotia loyolensis (Gill), and species of the strophomenid Leptaenisea, the leptaenid Leptagonia?, and allypidines including Spinall)pa (Jsospinallypa) sp. The sparsely OCCUlTing brachiopods from associated but younger limestone olistoliths oflate Pragian-earliest Emsian age (Mawson & Talent, 1994a) have not been described. A small brachiopod fauna of spiriferidine (HOlvellella?)

187

and rhynchonellidine brachiopods from the Lilydale Limestone (kindlei Zone; Wall et al., 1995) has yet to be documented.

The shelly fauna of the basal unit of the Wentworth Group, the Wild Horse Formation, in the watershed of the Mitchell and Wentworth Rivers includes poorly preserved, broken brachiopod material, but the occurrence of Polygnathus pireneae in limestone clasts from this unit provides a maximum age of late Pragian pireneae Zone for the base of the Wentworth Group and thus a maximum age for the faunas of the Dead Bull Member-possibly laterally equivalent to pali ofthe Wild Horse FOlmationand the Kilgower Member of the Tabberabbera Formation (Talent, 1963). The Dead Bull fauna includes the dalmanellidines Reeftonia mOlwicki (Allan) and Muriferella punctata (Talent), the spiriferidines "Adolfta" glypta Talent and Howellella aff. textilis Talent, and poorly preserved chonetidines andrhynchonellidines.

The Point Hibbs Limestone (sulcatus and kindlei zones according to Winchester-Seeto & Carey, 2000) at Point Hibbs on the W coast of Tasmania has produced an interesting brachiopod fauna (Flood, 1974). Only two identifications are currently accepted: Reeftonia manvieki (Allan) and Taimyn·hynx? globosus (Talent); there are species of allypidines and rhynchonellidines for which more information is needed by calcining, serial sectioning, and attention to microsculpture (Talent et al., 2000). Affinities appear to be with the Pragian faunas of central Victoria, e.g., the Boola Siltstone and Coopers Creek Limestone.

The GalTa Limestone at Wellington has more than 100 horizons of silicified limestones. Diverse brachiopod faunas (sulcatus Zone to inferred pireneae Zone, Wilson, 1989) have been monographed (Lenz & Johnson, 1985a, 1985b) Subsequently found late Lochkovian silicified faunas are presently being documented. The rich brachiopod faunas fi"Om the outcrop-tract on the E side of the Bell River (sulcatus and inferred kindlei zones) include the orthidines Dolerorthis ornata Lenz & Johnson and Ptyehopleurella forticostata Lenz & Johnson, the dalmanellidines Levenea mawsonae (Lenz & Johnson), Isorthis (Protocorlezorthis) impressiva Lenz & Johnson, DalejillCl ovata Lenz & Johnson, Schizo phoria la/isulcata Lenz & Johnson, S. fecunda Lenz & Johnson, Muriferella puncta/a Talent and Mys/rophora garraensis Lenz & Johnson, the chonetidine Asymmefroc!zoJletes planata Lenz & Johnson, the plectambonitidPleetodonta bipartita (Chapman), the stropheodontids Nadias/rophia superba Talent, Mesodouvillina (Protocymostrophia) magnifica (Lenz & Johnson) and Leptostrophiella (Rhytistrophia) leptocostata

188

Lenz & Johnson, the orthotetidine Eoschuchel'tella quadl'ata Lenz & Johnson, the rhynchonellidines Machaeraria catombalensis Strusz, Eoglossinotoechia catombalensis Lenz & Johnson and Sphaeril'hynchia globulal'is Talent, the altypidines AIiJ'pa invel'sa Savage,Atl)'Paria pellelopeae (Chatterton), PunctatlJ'pa (SinopUllcatlJ'pa) chattertoni (Lenz & Johnson), Spirigerina marginalijormis Alekseeva, and Coelospil'a septata Lenz & Johnson, the spiriferidines Cyrtill(l wellingtonensis Dun, Ambocoelia? dOl'siplicata Savage, Howellella australis Savage, H. latisulcata Talent, H. textilis Talent, Mauispil'ijel' hectol'i Allan, Howiftia howifti (Chapman), PlicocYl'tina coopel'i (Gill), Reticulariopsis talenti Lenz & Johnson and Quadl'ithYl'is tiro (Barrande), and forms not sufficiently well characterized yet of Skenidioides, Talentella, IsO/'tilis (Tyel'sella), Ctenochonetes, Chonetes (Plebejochonetes), Notoleptaena, Lepidoleptaena, Amphistrophia, Aesopomum, Pentamerella, Atrypina, Carinatina, Rugosatl)'pa, Australina, Plectospira, Kaplicona, Nucleospil'a, and Havlicelda.

Brachiopod faunas from the western tract of GatTa Limestone at Wellington (approximately pireneae Zone) are somewhat less diverse than those from the eastern tract (sulcatus-kindlei Zones). The faunas are similar but less diverse (14 species less), with the loss of Dolerorthis ornata Lenz & Johnson, Ptychopleul'ella jorticostata Lenz & Johnson, Levenea mawsonae (Lenz & Johnson), Mystrophora gal'l'aensis Lenz & Johnson, Plectodonta bipal'tita (Chapman), Leptostrophiella (Rhytistrophia) leptocostata Lenz & Johnson, Machaeraria catombalensis Strusz, AtJJpa inversa Savage, PUllctatlJ'pa (SillopUIICafl)'Pa) chatfertoni (Lenz & Johnson), Spirigerina marginalijol'mis Alekseeva, Howel!ella australis Savage, H. latisulcataTalent, Plicocyrtina cooped Gill, Reticulariopsis talenti Lenz & Johnson and the genera Ctellocholletes, Cholletes (Plebejochonetes), Notoleptaella, Amphistrophia, Aesopomum, Nucfeospira and Havlicelda. Noteworthy is the loss of what might be termed archaic elements: Dolerorthis, Ptychopleurella, Plecfodonta, Machaeraria, Spirigerina, Amphistrophia and Havlicekia, though not all of these may have gone into extinction between the kindlei and pireneae zones. There is a high level of endemicity at species level, but a remarkably low level of generic endemicity. Of the 47 genera represented, all can be descdbed as semi-cosmopolitan except two, Nadiastrophia and the subgenus Punctaflypa (Sinopullcatl)'Pa); these occur in S China as well as E Australia.

The Reefton Group of the Buller Terrane near Reefton on the W coast of the South Island of


New Zealand is now known from recent mapping (Bradshaw & Hegen, 1983; Bradshaw, 1989) to embrace several formations but allocation of the earlier documented brachiopod faunas (Allan, 1935, 1947; Boucot et al., 1963; Boucot & Johnson, 1967) to these units is not clear for several ofthe taxa. It is still uncertain how much if any of the sequence may be Pragian; it is conceivably substantially if not entirely Emsian, as was assumed by Talent et ai, (2000), The aggregate fauna consists of a dalmanellidine, Reejtonia maJ'wicki Allan, a strophomenid Maoristrophia neozelanica Allan, a stropheodontid Plicosfropheodonfa huttoni (Allan), a chonetidine, Allanetes neozelanica BOlicOt & Johnson, a rhynchonelIidine, Tanerhynchia parki (Allan), two spiriferidines, Mauispirijer itectori Allan and EUI)'spirijel' coxi (Allan), and two terebratulids, Reejionella lIeoze/allica (Allan) and Pleul'othyrella vellusta Boucot ef al. The presence of the otherwise Malvinokaffric Reaim Pleurothyrella is notewOlihy.

The pattern of linkages obtained from expeli systems analysis ofthe Pragian faunas ofthe AsiaAustralia hemisphere (Yolkina ef al., 1999; Talent et ai" 2000; Yolkin et ai" 2000) does not show pronounced biogeographic linkages during Pragian times between the Australian and New Zealand faunas on the one hand and contemporaneous faunas from the South China microcontinent, nor to other continental blocks in the Asian collage (Fig. 11). The regions of E Australia and New Zealand were neve.iheless shown to be biogeographically linked to each other during Pragian-Emsian times, the New Zealand faunas when compared with other regions of the Asia-Australia hemisphere tend to be biogeographically discrete due to presence of elements usually taken to be characteristic of the Malvinokaffric Realm, i.e. Antarctica, southern South America, Falkland Islands and southern Africa (see section on Bivalvia below).

EII/siall. Brachiopod faunas fi'om the Kilgower Member of the Tabberabbera FOlmation (Gill, 1949a; Talent, 1963; Johnson & Talent, 1967a, 1967b) are more diverse and inferred to be no older than pireneae Zone or perhaps as young as early in the dehiscells Zone, They are from a similar biofacies as the Boola and upper Humevale faunas and have a few species in connnon, notably the dalmanellidine Reejionia mal1vicki (Allen), the plectambonitidPlectodollta bipal'tita (Chapman) and the rhynchonellidine SphaerirhYllchia globularis Talent, but the majority of the species, despite being referrable to genera found in the Boola and upper Humevale faunas, are specifically distinct. These include the dalmanellidine Murijerella pUllctata Talent, the leptaenid


Rugoleptaella undulifera (Talent), the stropheodontids Nadiastrophia superba Talent, Leptostrophiella affillalata (Gill) and Cymostrophia bellarugosa Talent, chonetidines including Paracholle!es baragwanathi (Gill), the pentameridine Devonogypa? polymita (Gill), the rhynchonellidines Uncillulus calathiscus Talent and Eoglossinotoechia IOllgisepta Talent, the atrypidines Totia perflabellata (Talent), Spinatl)'Pa (Jsospinatl)'Pa) lIndosa (Talent), the spiriferidines Howellella piger (Talent), H.? pingllis Talent, "Adolfia" gl)'Pta Talent, Howellella textilis Talent, Plicocyrtina coo peri (Gill) andAltajella? grayi (Talent) together with less readily characterised species of Schizo phoria and other dahnanellidines, athylididines including Nucleospira and ?Buchanathyris, atrypidines including Australina? and spiriferidines including Cyrtina, several species of Hysterolites, and Eospirifer. The youngest unit of the Tabberabbera Formation, the Roaring Mag Member, has produced sheared athyrididines resembling Buchallathyris and delthyridids possibly congeneric with Spinel/a (Talent, 1963; Talent et al., 2000) implying a fauna of similar biofacies and perhaps broadly similar age to the Buchan Caves Limestone, but the low diversity and poor preservation prevent useful comment on biogeographic affinities.

Three faunas can be discriminated in the Buchan Group (pireneae to serotinus Zones, Mawson, 1987a; Mawson et al., 1988; Mawsonet al., 1992) ofE Victoria. The oldest, the low diversity fauna of the Buchan Caves Limestone (Talent, 1956a), is dominated by species of Spinella and Buchanathyris, with the spiriferidines S. buchanensis Talent and Howittia howitti (Chapman) appearing near the top of the Buchan Caves Limestone. This fauna is best known from the upper half of the fOlmation at Buchan, The Basin and Bindi and is early in the dehiscens Zone. Essentially the same fauna extends down into the lower patt of the formation to levels assumed to represent at least part ofthe pireneae Zone, though this has been demonstrated indubitably at only one locality, low in the Buchan Caves Limestone at The Basin, NE of Buchan. A smaller form of Spinella occurring at these levels may be S. yassensis (de Koninck) better known from the Cavan Formation and the lower units of the Taemas Limestone-to perhaps low in the Receptaculites Member'--Of the MUlnlmbidgee Group of the Taemas-Wee Jasper area of SE NSW.

The lower Taravale FOlmation, as typically exposed in the entrance cutting to the Buchan Caves Reserve, and the Pyramids Member ofthe Fonnation, are late dehiscens and early perbol111s

189

Zones. Salient brachiopods include the dalmanellidine Dalejina philipi (Chatterton), the chonetidines Parachonetes buchanensis (Gill), Protochonetes auslralis (McCoy), the atlypidines Coelospira dayi Chatterton and Desquamalia (Variah)'Pa) erectirosl1'ts (Mitchell & Dun), the athyridines Athyris waratahensis (Talent) and Buchanathyris westoni Talent, and the spiriferidines Ambocoelia? rll"negari (Chatterton), Quadrithyrina allani Chatterton, Howittia howitti (Chapman), rare H. muitiplicata (de Koninck) and Spinella buchanensis. The chonetidine Septachonetes teicherti (Gill) is abundant in nodules at some horizons high in the Pyramids Member. Brachiopods are VCly rare in the uppermost levels of the Taravale Formationlatest perbonus, inverSllS and sera/inlls Zones(Mawson 1987a; Mawsonel 01.,1992).

The Murrindal Limestone faunas (late dehiscens and perbonus Zones; Mawson, 1987a and unpub. data) are more diverse (Talent, unpub. data); some of the horizons are silicified. The faunas include many elements of the lower Taravale-Pyramids fauna, namely the dalmanellidineDalejina philipi (Chattelton), the chonetidinesParachonetes buchanensis (Gill) and Protochonetes auslralis (McCoy), several atrypidines including Coelospira dayi Chatterton, Desquamatia (Variallypa) erectirostris (Mitchell & Dun), and the spiriferidines Howittia howitfi (Chapman), H. multiplicata (de Koninck) and Quadrithyrina allani Chatterton. Among other elements are the generically endemic stropheodontid Malurostrophia flabellicauda Campbell & Talent-abundant in some hOlizonsand Mesodouvillina (ProtoCYlllostrophia) dickinsi (Chatterton), the orthotetine Eoschuchertella /tIlI/phyi (Chattelton), rhynchone11idines including Eoglossinotoecilia? linki Chattelton and several undescribed forms, several attypidines including Atlyparia penelopeae (Chatterton) and several undescribed spirifelidines. The faunas have much in common with the approximately coeval faunas of the Receptaculiles and Warroo Limestone Melllber~ ofthe Taemas Limestone, Taemas, NSW (Chatterton, 1973).

Brachiopod faunas from the Cavan and Majurgong Formations and of the "Spirifer" yassensis, CUlTajong and Bloomfield Limestone Members of the Taemas Limestone ofthe TaemasWee Jasper area ofNSW have low diversity and have not received much attention. Most attention has been given to the "Spirifer" yassensis Limestone Member, a unit with a low diversity fauna (Strusz el al., 1970; Chattelton, 1973; cf. Talent et aJ., 2000) very similar to that of the Buchan Caves Limestone: Protoclionetes aush'alis (McCoy), Athyris waralahensis (Talent),

190

Spinella yassensis (de Koninck), Howittia howitti (Chapman) and "H. " multiplicata (de Koninck)

Diverse early Emsian (perbollus Zone) silicified brachiopod faunas (e.g., Fig. 10) have been monographed from higher units of the Taemas Limestone (Stmsz etal., 1970; Chatterton, 1973; cf. Talent et al., 2000). The most diverse fauna is from the Receptaculites Member. It consists of Salopina kemezysi Chattelton, Dalejina philipi (Chatterton), Mesodouvillina (Protocymostrophia) dickinsi (Chattelton), Malurostrophia flabellicauda Campbell & Talent, Mesoleptostrophia (Paraleptostrophia) clarkei (Chatterton), Eoschuchertella mllIphyi (Chatterton), Parachonetes buchanensis (Gill), Protochonetes australis (McCoy), Chattertonia campbelli (Chatterton), BrOlvneella browneae Chattel1on, Eoglossinotoechia? linki Chatterton, Coelospira dayi Chatterton, AtlJ'paria penelopeae (Chatterton), Desquamatia (Variaflypa) erectirostris (Mitchell & Dun), Athyris warafahensis (Talent), Ambocoelia? rUl1negari (Chatterton), "Delthyris" hudsoni Chatterton, HOlVittia howitti (Chapman), H. multiplicata (de Koninck), Spinella yassensis (de Koninck), Quadrithyrilla allani Chatterton, Plicocyrtina crenulata Gratsianova & Talent,Adrenia expansa Chattel1on, Cydimia roberts; Chattelton, Micidus shalldkyddi Chatterton and M? glaber Chatterton, and unidentified Cyrtina.

A somewhat less diverse silicified brachiopod fauna has been obtained :liOln the overlying Warroo Limestone Member (pel'bOllUS-inversus zones) of the Taemas Limestone (Chattelton, 1973; Talent et al., 2000) consisting of Isorthis (Tyersella) spedelli (Chatterton), Resserella careyi Chatterton, Muriferella punctata (Talent), Dalejina philipi (Chatterton), Mesodouvillina (Protocymostrophia) dickinsi (Chatterton), Nadiastraphia patmorei (Chatterton), Malurostrophiajlabellicauda Campbell & Talent, Mesoleptostrophia (Paraleptostrophia) clarkei (Chatterton), Eoschuchertella murphyi (Chatterton), Protochonetes australis (McCoy), Parachonetes buclwllensis (Gill), Septachonetes teicherti (Gill), Chattertonia campbelli (Chatterton), Eoglossinotoechia? lillki Chatterton, Coelospira dayi Chatterton, Ambocoelia rUl1l1egari (Chatterton), Howittia howitti (Chapman), 'Delthyris' hudsolli Chatterton, Adrenia "-'pansa Chatterton, Micidus shandkyddi Chattelton and species of Cyrtina. Similar silicified faunal assemblages are known from the Brogans Creek Limestone (Colquhoun, 1998) and from low in the Mount Frome Limestone of the Mudgee district (A.J. Wright, pel's. comm.). Higher in the Mount Frome Limestone are silicified Megastrophia and Zdimir (A.J. Wright, pers.


comm.); this occunence is consistent with the Emsian-Eifelian boundary OCCUlTing near the top of that formation (cf. Pickett, 1978; Mawson, 1987b, p. 256).

Conodont data from fossiliferous sequences of the often poorly outcropping Ukalunda Beds on Mary Creek and Douglas Creek in NE Queensland, about 200 and 400 km respectively S of Townsville, are indicative of the early Emsian perbollus Zone. The brachiopod faunas fi'om these and a third locality are essentially the same as, though less diverse than those of the Receptaculites Limestone Member ofthe Taemas Limestone of SE NSW (Parfrey, 1989; Brock & Talent, 1993): IsO/·this (Tyersella) spedeni (Chatterton), Aulacella philipi Chatterton, Mesodouvillina (Protocymostrophia) cf. dickinsi (Chatterton), Nadiastrophia patmorei (Chatterton), Mesoleptostrophia (Paraleptostrophia) clarkei (Chatterton). Atryparia penelopeae (Chatterton), VilriatlJpa (VariaflJpa) erectirosMs (Mitchell & Dun) and Carinatina lowtherensis Johnson & Boucot as well as undescribed species of Hypsomyonia, Parapugnax, Quadrithyrina, Warrenella and Elythina. The great species-level similarity between these faunas from about 1300 km apart palaeolongitudinally is consistent with little or no major translation of the respective crustal blocks with respect to each other since early Emsian times.

Middle Devonian, Eifelian brachiopods are known from the Sedgeford Formation at Alpha, central Queensland (Henderson et al., 1995) and from an earliest Givetian (hemiansatlls Zone) horizon in the Moore Creek Limestone at 'WalTawilla' near Attunga, NE NSW-where they are silicified. A possible Eifelian fauna has been described from the Etonvale FOlmation in the subsUlface Adavale Basin ofSW Queensland'. Bornhardtilla coulteri has been documented (Brown, 1943) from a horizon close to the Eifelian-Givetian boundary in the Moore Creek Limestone at Sulcar, a few km N of Attunga, but this form is a species inquerendum. Stringocephalus (identified as S. burtiniDefrance by Brown, 1943) and Warrenella occur at many localities in the Burdekin Limestone, e.g., near 'Fanning Downs' and on the Reid River, but have not been described. The overlying late Givetian Cultivation Gully Formation has a diverse fauna (McKellar, I 967}-in the Limestone Creek area S of Fanning Downs-with the spiriferidines Cyrtinaella and Warren ella, schuchertellids, rhynchonellids, and atrypids such asDesquamatia (NeaflJpa), but the fauna has not been described. The only adequately described occurrence of the typically early Givetian centronellidine Stringocephalus is S. fOlltanus from the Pillara


Limestone at Mountain Home Spring on the nOlihern flank of the Canning Basin of Westem Australia (Veevers, 1959a),

The most diverse Middle Devonian brachiopod faunas from Australia occur in the Papilio Formation (Givetian to at least as young as hermanlli Zone) of the Broken River region of the Townsville hinterland, NE Queensland, The faunas are noteworthy for high diversity of atrypidines but velY few f011115 have been described (Brock, 1989; Gratsianova et al., 1990): Desqllamatia (Desquamatia) peshiellsis (Grabau), Desqllamatia (Illdependaflypa) maglla (Grabau), D. (1.) zonatoides Biernat, Desquamatia (SyJlGflJ'pa) subzonata (Bielnat), Kelpina vineta villeta Suuve, Spillallypina (Spillaflypilla) sO'eblo (Chen), Grllellewaldtia latilingllis (Schnur), Carinatina plana (Kayser), Davidsonia verneuili Bouchard-Chantereaux, Warrenella dorothae Talent and species of Atl)'Pa (Kyrtatrypa), Atlyparia, Variatlypa (Variatrypa) and Spinah)'Pa. The faunas are notewOlihy not only for their generic and specific diversity, but for the number of species conspecific with fOlIDS known from elsewhere in the Old World Realm. A meaningful statement on biogeographic affinities must however await documentation of the entire fauna. It is not clear ifthis apparently low level of endemicity extends throughout these diverse faunas.

Late Devonian, The Frasnian brachiopod faunas of the Canning Basin (Coleman, 1951; Veevers, 1959a, 1959b, 1959c; Sartenaer, 1970, 1971, 1972, 1979; Grey, 1978; Strusz, 1992) are noteworthy for prominence of atrypidines, all seemingly endemic at the species-level: Desqllamatia (Synatlypa) kimberleyensis (Coleman) in the Gogo Formation; D. (S.) kimberleyensis and Spillall)'pilla (Exatr)'Pa) kuniandia Grey in the shallow platform Pillara Limestone; and the same two forms together with Atrypa (Kyrtatrypa) teicherti (Coleman), Spinaflypilla (Spillatrypina) pride!'i pride"; (Coleman) and S. (S.) prideri Illlrllllgunia Grey in the Sadler Limestone. Also prominent are the rhynchonellidines Ptychomaletoechia Ilicida (Veevers), Fitzroyella primula Veevers and PhlogoiderhYllchlis are/actus (Veevers) from the Sadler Limestone-with the last occurring in the Napier Limestone as well-and Hypseloterorhync/tus pennatus Sartenaer, Parvulaltarostrum veeversi Smienaer and Cavatisinurostrum sp. in the Virgin Hills Formation, and Flabelllllirostrum wolmericum (Veevers) occurring in several early Frasnian units. A sequence of seven macrofaunal "zones" was proposed by Veevers (1959a), all except one of

191

them based on brachiopods. The relative ranges of brachiopods with respect to these "zones" are presented in Table 2, but how these align with the conodont zonal schemes is imprecise. Noteworthy is that there was a considerable decline in diversity during the Frasnian long before the Kellwasser events; the reason for this is probematic. More data are necessalY before one can be sure whether or not this was connected with a global event or events. Of the Frasnian fomls, all species appear to be endemic and about 25% of the genera appeal' to be endemic, this runs counter to the widespread assumption that Frasnian brachiopod faunas were fundamentally semicosmopolitan (cf. the situation with regard to Frasnian conodonts). The Famennian brachiopod faunas were less diverse and, at generic level, more cosmopolitan than the Frasnian.

In the Bonaparte Gulf Basin ofNW Australia, three members around the middle of the Cockatoo Formation-the Kununurra, Hargreaves and Westwood Members-have Frasnian brachiopod faunas consisting, in aggregate, of Productella lvestwoodensis Roberts, Retichonetes arenarius Roberts, Globosocholletes? rnathesonellsis Roberts, Spinatlypina (Spinatlypina) prideri larga (Roberts), Desquamatia (SYllatITpa) kimberleyensis (Coleman), CrurUhyris apena Veevers and Tenticospirijer columnaris Robelts together with unidentified or indetelminate fonns of Schuchertella, Sieinhagella, Calvillaria and Cyrlospirijer (Roberts, 1971). The age indicated by conodonts is approximately the interval represented by the former Ancyrognathus h'iangularis to Early gigas zones (Robelis et al., 1972), i.e. about zones 11 and 12 of Klapper (1989; cf. Klapper & Becker, 1999) or about jamiae Zone to the early part ofthe rhenana Zone of Ziegler & Sandberg (1990). The age is imprecise because the conodonts are from icriodid rather than pahnatolepid biofacies. Despite proximity to the Canning Basin, only C. apena, a zonal form in the latter, provides any sort of biostratigraphic alignment on the basis of brachiopods. All genera represented are cosmopolitan but, at species-level, endemism is high.

The reefal Ningbing Limestone, overlying the Cockatoo Formation of the Bonaparte Basin, spans the Devonian-Carboniferous boundary. The Late Devonian (Famennian, rhomboidea Zone and later) brachiopod faunas ofthis unit consist ofthe productidines Sentosia subqlladrata Roberts and Mesoplica jeremiahensis Roberts, the rhynchonellidines Ptychomaletoechia lucida (Veevers), RlIgaltoroso'ulII australe Roberts and Nayunella turgida Robelis, and the spiriferidine Cyrtospirijer ningbingensis Roberts, together with unidentified or indeterminate species of

192 AAP Memoir 23 (2000)

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SlringocephaTusfonldJllIS Veevers • Rcticulariopsis 5IU:MOO (Veavers) • D<"""",.6:iJ (SY""'r:YP"J /ci,.,-hri<J<r.ji< (Coleman) • • • • Gypidulafragilis Veevers • If)p;:omyonia nip!taJw Veevers • liuljia sa/liea Veevsrs • Productella? ocdduG Veevers • Skenidiwn (lul/atum Veavers • Tfichertina. filuoycmis Veevers • Zophmtrophfa Wlgamica Veavers • \ ~<l"i'f>alooia nwnida (Veevsrs) • Sclmchemlla 8fl!lillic(l Veevers • CyphopleroriQnch,,5 puteanus (Veevers) • SpiF.atrypirdJ. (EmiTypa) hmimulia Grey • Spina.If)P,l/Spinalr),pa) mm11lgWlia Grey • Hm::mirophia nqllisila (Veevers) • • SchizopJwria stainbn..'oki Veevers • • 0

~ !< Spir<i1/0pa (Kyrtatrypa) tdcheni (Coleman) • • Dmp/arMtia (NearrJpa) multimoda Grey • • D«'onoproductrrs aIJStraJis Veevers • • FilZroyelfa prirnllla Veevers • • Flabdlllliroslmm wolmerkum (Veevers) • • Hypolhyridma margarita Veevers • • Nen"oslrophia butU1pica Veevers • • • Athyris oscarmsis Veavers • • Emanuella torrida Veevers • KtI)urdla eJr>111111eiem;is Veevers • P/ico<:honms IIWCWpalUS Vaevars • Spir.utrypir ... (Spir.ulrypir-<» pri&rl pOOri (Co!MWlj. • P/r1ogoider/rj7!chus arifactus (Veevers) • • Cnuithyris apena Veevers • POf\,,/o/laroslrum ruwrsi Sartenaer • Hypselol<rorIrJ7!cJIIlS [k7lJ!.afIlS Sartenaar • !'.'Jose scopimlLs Veavers • Pug1UlX hullmsis Veevers • Schizophoriapiem,mds Veevers • Maislella? caprirw Veevers • • Niguillop/ira pn>ieus (Veevers) • PtychoffUJ/etoechia lucidn (Veevers) • Rhipidomdfil lncompln Veevers • Sch;zophoria apiwl<lla Veevars • Schllchertella dromeJo Vaavers •

AAP Memoir 23 (2000) 193

Fig. 11. A selection of species representative of Early Devonian (early Emsian) brachiopod genera from the early Emsian Yukiang Fonnation (coeval with faunas from the Buchan and Taemas Groups ofSEAustralia), believed from available data to be endemic to the South China Province or restricted to few provinces:A-C,Dicoe/ostrophia crenafa Wang. Exterior, interior and anterio[views respectively of pedicle valve, xl.S. D,PullctallJpa (Undahypa) bellalllla \Vang ef al. Exterior view of pedicle valve, x5. E, F, HOlvittia modica \Vang & Rang. Anterior and lateral views respectively of complete specimen, x2. G, Barbathyris glabra \Vang & Rang. Illustration of interior of brachial valve showing brachidium, x4. Drawings by Julie Trotter Ii-om Wang & Rong (1986).

Schllchertella and Prodllctella (Roberts, 1971). Again, the genera represented are cosmopolitan but at species-level, there is 100% endemism.

The brachiopods of the Gneudna Formation of the Carnarvon Basin of West em Australia include a productidine, Productella? oecidua Veevers, a rhynchonellidine, Cyphopterorhynchus pl/leana (Veevers), the widely distributed atrypidine, Spinallypina (Spinaf1ypina) prideri (Coleman) and five species of spiriferidines: AlIslrospirifer variabilis Glenister, CyrtospiriJer minilyaensis Glenister, C. australis Glenister, C. gneudnaensis Glenister and C. brevicardinis Glenister (Glenister, 1955; Veevers, 1959b)

There has been no work on the Late Devonian brachiopod faunas ofE Australia in the past thirty years. The review by Roberts el al. (1972, p. 478-480) is thus still a useful survey of these faunas and correlations in the Late Devonian for E Australia'. Generally poorly preserved, low diversity brachiopod faunas often dominated by Cyrlospirifer with subordinate rhynchoneJlidines,

occasional atrypidines (exclusively Frasnian), and productidines, the last becoming more prominent in Famennian horizons, have beenrepOlied fi'om many areas in E Australia from Eden in SE NSW to the Yarrol 'Basin' of central Queensland (specifically the Monto-Mount MorganRockhampton area) and the Mt Wyatt area and Burdekin Basin ('Star Basin') of the Townsville hinterland (Maxwell, 1951, 1954, McKay, 1964; Robelts el al., 1972). The taxonomic position of the dominant cYltospiriferids relative to the vast number of nominal species of Cyrtospirifer (or synonyms of CyrtospirifelJ, Tenticospirifer and closely related genera globally is problematic. McKellar's (1970) work on the productidines of the Burdekin Basin resulted in a series of zones; these have proved useful for approximate correlations ofthe Frasnian-Famennian sequences of the Monto-Mount Morgan-Rockhampton area. Prominent in the Famennian faunas of the Burdekin Basin are species of the productidines SemiplVductus, Sentosia, Spillulicosta, Laminatia

Table 2 (opposite). Distribution of Camting Basin Devonian brachiopods (Coleman, 1951; Veevers, 1959a, 1959b, 1959c; Sartenaer, 1970, 1971, 1972, 1979; Grey, 1978) based on analyses by Grey (1984a, 1984b) and Stmsz (1992). Alignment of the brachiopod 'zones' with the conodont zones is highly approximate. Only taxa identified to species-level are shown. Note that most of the described taxa are from horizons equating with the interval covered by the early Frasnianfalsiovalis-punctata zones, appreciably pre-dating the Lower and Upper Kellwasser Events. Note also the lack of a significant database through the latter global extinction events.

194 AAP Memoir 23 (2000)

East Russian

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Fig. 12. Minimum spanning tree (MST) based on Jaccard coefficients for the 17 regions from which significant Pragian brachiopod faunas have been documented, showing names of regions and the Jaccard similarities between them (from Yolkin et a/" 2000).

and Mesoplica? These genera are relatively cosmopolitan and, though all species were described as new, closest affinities of the various species are with fonns scattered across Laurentia and Laurussia. The biogeographic significance of these faunas thus remains problematic.

COlle/usiollS. The presently available database for the Early to Middle Devonian brachiopods of the Asia-Australia hemisphere has been revised in order to obtain taxonomic consistency at generic and species level (Talent et aI., 2000) but, to date, has been subjected to computer-analysis only for the Pragian interval. This has demonstrated there were strong linkages between the brachiopod faunas of the various tenanes ofSEAustralia and New Zealand, and that these linkages were stronger than with contemporaneous brachiopod faunas from the crustal blocks of Asia (Yolkin et ai., 2000), including South China (cf. Figs 10 and 11; Fig. 12). The latter is juxtaposed with N Australia in several reconstructions for Early Devonian time. If this had indeed been the configuration, a higher level of similarity would be expected. Viewed qualitatively, the same lack of similarity is the case for the Lochkovian and

early Emisian interval. There are, nevertheless, contrasts between the

Early Devonian brachiopod faunas from tenane to tenane in E Australia. These contrasts may be a reflection of profound differences in

sedimentalY environments, especially clastics compared with carbonate environments, but could include effects of substantial displacement causing juxtaposition of telTanes that may once have been much farther apart (Talent, 1985b). This cannot be conclusively determined without quantitative biogeographic analysis based on a more substantial taxonomic database than is presently available.

The New Zealand Early Devonian faunas, specifically the faunas ofthe Reefton Group, are more provincial than one might anticipate from casual examination (Yolkin et ai., 2000); they have an interesting Malvinokaffric component compared with SE Australia.

Australian Middle Devonian brachiopod faunas are still too poorly known for meaningful evaluation of biogeographic linkages.

Species-level endemism is high for the Late Devonian brachiopod faunas of West em Australia. This is unexpected in view of the cosmopolitan nature of the coral faunas (A.J. Wright, pers. comm.). Generic·level provinciality seems to have been absent for the Bonaparte Gulf Basin (though the faunas are not velY diverse), but was appreciable at generic level-up to 25%--in the case of the Canning Basin.

Algae (J. Douglas, J.A. Talent & R. Mawson) As in the Silurian, non-calcareous algae seem

to have proliferated in the Early and Middle


Devonian seas of E Australia. Though occasionally occUlTing in re1ative abundance at specific localities in flyschoid enviromnents, their preservation as generally tenuous carbonaceous films makes them taxonomically nnattractive. Prominent among such algae and questionable algae are forms - following Lucas (1927) and, before him, Chapman (1903) - referred to Blithotrephis Hall, a name used to embrace a multitude of putative and indeterminate fossils. A half-dozen species of marine algae, based on better preserved material, have now been described (Douglas & Jell, 1985) including Blithotrepizis trichotoma Douglas & Jell from the Humevale Siltstone (Lochkovian-?Pragian) and B. walhal/a Douglas & Jell from the Norton Gully Sandstone, Walhalla (Pragian) of central Victoria. The present database is insufficient as a basis for hazarding comment on biogeographic affinities.

Calcareous algae are prominent in the numerous tracts of shoalwater and shallow marine limestones in the Devonian ofE Australia. Such floras, for the most part, remain to be documented, but important contributions have been made on calcareous algae from the Early Devonian (principally late Lochkovian-Pragian) limestone olistoliths in the Nubrigyn Member of the Cunningham FOlmation (late Lochkovian pesavis Zone to late Emsian; Talent & Mawson, 1999) of the Stuart Town-Euchareena area of east-central NSW. In 1964 J. Harlan Johnson described 13 species of algae from these olistoliths (thought at the time to be autochthonous); one genus and 7 species were described as new. His conclusion that these floras displayed shnilarity to algal floras from the Kuznetsk Basin ofSW Siberia (Maslov, 1956) must be considered in the context of the limited database on Devonian calcareous algae at that thne. Also of latest Lochkovian-Pragian age are undescribed, diverse algal floras with prominent Lancicula from the Arch Creek and Martins Well Members of the Shield Creek Formation of the Broken River region of NE Queensland. Algae are a major component of shallow marine limestones in the Tamworth Belt ofNE NSW, in late Emsian Sulcor and Eifelianearly Givetian Moore Creek and Timor Limestones. For example, from an algal limestone horizon within the Moore Creek Limestone at 'Wyaralong', Attunga, Pohler & Herbert (1993) report the presence of species of Girvanella, COllvinianella, Sphaerocodium, Renalcis and possibly Peisteignella.

Late Devonian floras of the Canning Basin were documented by Wray (1967). Of the 12 genera and 16 species documented, 3 genera and 11 species were recognised as new. Genera represented included red algae (Solenopora,

195

Parachaetetes, Stenophyclls, Keega, Tharama), green algae (Litanaia, Ortonella, Vermoporella), with cyanophytes (Girvanella, Sphaerocodium, Renalcis, Paraepiphyton) being especially abundant and diverse. Algal floras of the Late Devonian limestones of eastemAustralia have not been documented, though Renalcis is prominent in limestone clasts in the Mostyn Vale Formation ofNE NSW (A.J. Wright, pers. comm.).

Numerous other algal floras remain undocumented, notably from the LochkovianPragian Garra Limestone of east-central NSW, the early Emsian Murrindal Limestone ofE Victoria, the limestone units of the Broken River Group (late Emsian-earliest Frasnian?) of the Broken Riverregion ofNE Queensland, and the Mt Podge Limestone (late Emsian), Burdekin Limestone (late Eifelian-early Givetian) and the lenticular limestones in the Myrtlevale Formation (midFamennian) of the Burdekin Basin of NE Queensland. Until monographic studies of the Devonian algal floras, paliiculariy ofEAustralia, have been undertaken, biogeographic implications of the Australian algal floras must remain unclear.

Radiolaria (J.C. Aitchison) Data are presently insufficient to permit clear

discrimination of provinciality among the relatively few occunences of AustralianDevonian radiolarians, but radiolarian assemblages recently proposed for the late Early and Middle Devonian ofE Australia are not easily discliminated globally (Stratford & Aitchison, 1997; Aitchison et al., 1999)10. Substantial endemism is thus apparent. The taxonomically most diverse and bestpreserved Devonian radiolarian faunas in Australia are from early Frasnian carbonate concretions in the Gogo Fonnation of Western Australia (Nazarov ef al., 1982; Nazarov & Ormiston, 1983; Aitchison 1993b). These faunas display affmities with coeval radiolarian faunas from North America, the S Urals and ByelolUssia and may therefore be viewed as indicating greater ease of communication than had been the case earlier in the Devonian.

Foraminiferida (K.N. Bell) Few studies on Australian Early or Middle

Palaeozoic foraminiferal faunas have been undertaken on a regional scale, most previous studies being confined to a specific sequence or to relatively close sample sites. Information regarding facies dependence of foraminiferal faunas during these times is thus not well understood. Identification of very simple foraminiferal forms - e.g., species of Psammosphaera, Rhabdammina and Sorosphaera - is

196

often problematic; many ofthe described species may be synonyms. Many stndies on Early or Middle Palaeozoic foraminiferal faunas from elsewhere in the world tend to provide only broad ages and very generalised information on the horizons sampled. Some biogeographic patterns, albeit broad, can nevertheless be discerned.

Australian Lochkovian and Pragian foraminiferal faunas (Bell, 1999; Bell ef al., 2000) display about 40% endemicity at species-level. By Emsian times there was much higher endemism-at least 60% (Bell, 1996). This high level of endemicity appears to be valid, given the extensive documentation of contemporaneous foraminiferal faunas from the Emsian of the USN°. Endemism appears to have decreased in the Eifelian with a drop to 50% endemic species (Bell & Winchester-Seeto, 1999)11.

Few data are available on Late Devonian foraminiferal faunas from Australia (Crespin, 1961; Conkin & Conkin, 1968) but foUl' genera and eight species described from the Canning Basin by Conkin & Conkin were deemed to show strong similarities to faunas fi'om USA and Europe (Conkin & Conkin, 1968) thus suggesting a low level of endemism during this time interval.

Porifera (J.W. Pickett) Two relevant publications (Rigby, 1986;

Pickett & Pohler; 1993) have appeared since a fairly recent overview ofthe literature concerning Australian fossil sponges (Pickett, 1983). Previous comments on paJaeobiogeographic affinities of Australian fossil sponges have been brief(Pickett in Talent, 1972; Rigby, 1979, 1986; Pickett & Pohler, 1993).

Sixty species of sponges have beenrepOlted from the Australian Devonian; 43 of them have been regarded as endemic. The apparent endemicity is high-72%, but several of the supposedly nonendemic taxa have been identified only to categories higher than genus, so the true level of endemicity may be appreciably higher. Of the 33 genera repOlted, 25 (76%) are endemic. The eight nonendemic genera include the widespreadHaplistiol1, Anfilaspidella, Hyalosfelia andAstyiospongia. The occurrence of Asty/ospongia tarda in the Lochkovian part of the Garra Formation is the youngest known repOlt of the genus. Occlmence of the sphinctozoan Hormospongia in the Middle Devonian (Bifelian) ofE Australia and Alaska has been interpreted (Pickett & PohleI', 1993) as reflecting a proto-Pacific connection. This recalls linkages indicated by conodonts (see elsewhere). Pelicaspongia in the Late Devonian reef assemblages of Western Australia accords with a degree of linkage-unexpected-with E North American faunas at that time.


As is common in mid Palaeozoic strata, the assemblages are dominated by anthaspidellids. The next most abundant group is the Astylospongiidae. This is unusual, as already pointed out by Rigby (1986, p. 36), who regarded their abundance as a diagnostic featnre of the Australian faunas. Perhaps the most distinctively Australian group consists offonns referred to the Columellae-spongiidae by Pickett (1969). None of the three genera (Coiumellaespongia, CrGlvneya, Oremo) has so far been reported outside Australia. The type genus was widespread, occUlTing at several localities in NSW and in NE Queensland; their large size and distinctive morphology make for easy recognition and identification.

Stromatoporoidea (B.D. Webby & Y.Y. Zhen) Stromatoporoids are distributed widely in the

Devonian carbonate-dominated successions of Australasia12• These sessile benthic organisms of problematic calcified sponge affinity lived in warm, shallow, well-agitated, marine environments, typically free of associated tel1'igenous material. They are significant framebuilding contributors of organic buildups (biostromes and/or bioherms), occurring in associations with bahamitic-type carbonates in the Early-Middle Devonian of E Australia, in the Early Devonian of New Zealand, and in the Late Devonian of the Canning, Carnarvon and Bonaparte Gulfbasins ofW Australia.

Devonian stromatoporoids have been widely regarded as indicators of shallow marine, tropical to sUbtropical environments, between 400N and 35°S (Heckel & Witzke, 1979), though the recent global survey of Stock (1990) shows a wider latitudinal spread, up to 600 N and 45'S. The palaeogeographic map bases of Scotese (1986) used by Stock (1990) may, however, be in need of revision given that, in one single lithospheric plate (Gondwana), the closest Devonian coral! stromatoporoid reefs to the Devonian south pole are in NW Africa, approximately 55' of palaeolatitude away from the pole (i.e., the reefs are placed at 35°S palaeolatitude), a scenario more in accord with Heckel & Witzke's estimates.

Stock (1990) repOlted a progressive decline in generic-level provincialism through Devonian time, initially between the Old World and the Eastern American (Appohimchi) realms tln'ough the Early Devonian, to a maximum of25% genera in common by the Emsian, then rising to 75-85% of genera in common during the Middle Devonian, even higher in the Late Devonian (Frasnian). No stromatoporoids have been recorded from the cooler Malvinokaffric Realm. The Australasian faunas have been assigned to


the Old World Realm, together with those of Eurasia, N Africa, western N America and the Canadian Arctic.

Thirty-nine stromatoporoid genera have been documented from the Devonian of Australia, but are not unifonnly distributed through the column. Only one small fauna has been repOlted from the Lochkovian: from the Elmside Fonnation of the Yass area, NSW (Birkhead, 1978); it has little biogeographic significance.

Pragian and Emsian faunas provide more useful insights into Australasian biogeographic trends. Several genera have specific restricted provincial relationships. Pattems based on the assembled global distribution data in Steam et al. (in press) help to clarifY the most likely linkages between the Australasian assemblages and those of neighbouring regions. Shared occunences are listed below for several Pragian-Emsian genera. These are the c1athrodictyid genera Schistodictyoll (Victmia, N Queensland and Kuznetsk Basin) and Atelodictyon (Victoria, N Queensland, Kuznetsk Basin, NE Siberia and Tien Shan), the actinostromatid genusPlectosD'Oma (Victoria, NE Siberia and Uzbekistan), the stromatoporellid genus Tubuliporella (with related species in Victoria and the Altai Mountains), the stromatoporoid genus Salairella (Victoria, NSW, Arctic Canada, Salair and the Omulevsk Mountains of NE Siberia), and the syringostromatid genus Atopostrorna (Victoria, NSW, S China,Arctic Canada, and a slightly later, Eifelian, record from the Kuznetsk Basin). Also of importance are genera such as the clathrodictyid Tienodictyon that shows changes from a restricted link (N Queensland and NE Siberia) to wider connections (Queensland, S China, NW Territories, E Urals, Kuznetsk Basin and Salair) through the Early to Middle Devonian interval, and the stromatoporid genus Pseudotrupetostroma that has a restricted NSW and Victor.ian record in the Emsian, but spreads to the Kuznetsk Basin and the E Urals during the Eifelian. Similar biogeographic trends emerge from study of species relationships. A few Pragian species from the Lilydale Limestone and cOlTelatives in central Victoria show close Asian affinities, for example, Syringostromella zilllchenkovi (Khalfma) is also knownfr'om Salair, and Plectostroma aitum (Ripper) has close relationships with other species of Plectostroma recorded from the Kuznetsk Basin and the Ancennis Basin (France).

Emsian species from the Jesse Limestone at Limekilns, central NSW, also show close provincial links (Webby & Zhen, 1993). AtoposD'Oma dis tails (Ripper) occurs also in the early Emsian Buchan Caves Limestone, E

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Victoria, as well as in Arctic Canada and the NW Territories. The type species of the genus, A. tUlltouense Yang & Dong from S China, is a closely related form.

Salairella prima Khromykh is another fmill with occurrences in Arctic Canada, and NE Siberia (Omulevsk Mountains), and Pseudotrupetostromajessiense Webby & Zhen is related to the type species P. artyschtensis (Yavorsky) from the Kuznetsk Basin. Emsian species from the Broken River region of N Queensland inclnde Atelodictyon cf. lazutkini (Yavorsky), again with possible links to the Kuznetsk Basin.

The early Eifelian record in N Queensland includesPseudoactinodictyon lozveuse Yavorsky, previously described fi'om the Urals. Another link is indicated by the occunences of Petridiostroma clarum (Pocta) in the early Emsian Buchan and Munindallimestones of E Victoria, the Middle Devonian Koneprusy Limestone of the Czech Republic, and possibly also from the Reefton succession of New Zealand. Nexililamina is the only endemic genus recognized in the late Emsian-Eifelian assemblages ofthe Broken River region.

The succeeding Givetian-Frasnian faunas, based on described occunences in N Queensland, and the Canning, Cammvon and Bonaparte basins ofW Australia, are much less provincially distinct. They are characteristically abundant in reef complexes and exhibit culmination of the trend to a cosmopolitan fauna. Twenty-six species have been reported by Cockbain (1985) from the Frasnian of the Canning Basin. Only one genus (ElIlJ'amphipora) appears to have a restricted distribution (Canning Basin, Alberta, Saskatchewan, Afghanistan). Otherwise most of the fannas are related (conspecific) with previously described European andlor N American forms, especially the BelgianArdennes faunas documented by Lecompte (1951,1952).

The dramatic reduction in diversity and abundance across the Frasnian-Famennian boundary led to a relict stock of labechiids, stromatoporellids and stromatoporids (only five remaining species) in the W Australia basins (Cockbain, 1989). Only one biogeographic ally significant feature is indicated by this Famennian fauna, namely the common appearance of labechiids in the Bonaparte Gulf Basin and S China - two identical late Famennian (,StlUnian') species, Penllastl'oma yangi Dong and Platijel'osh'Onta kueichowense (Dong), once more establishing the strong direct Australia-E Asian link.

In summary, the distribution patterns of Australasian stromatoporoid faunas accord with

198

suggest strongest levels of provincialism in the Early Devonian (Pragian-Emsian), followed by steady decline in provincialism to the Frasnian. The charactedstic aft1nities ofthese faunas is with Asia and Arctic N America, in particular with the Kuznetsk Basin of Siberia and other Asiatic Palts of thc fOlmer Soviet Union, as well as with S China, and the Canadian Arctic and NW TetTitories. A few faunal elements exhibit clearcut trends towards less restricted (wider) distribution patterns tlu'ough the Early-Middle Devonian transition, towards more cosmopolitan, circum-equatotial, "reef' assemblages of Givetian to Late Devonian (Frasnian) times.

Rugosa (A.J. Wright & Y.Y. Zhen) Australasian Devonian rugose corals are

known fi'om three regions: Early, Middle and Late Devonian faunas from the Tasman Fold Belt of eastern Australia; Givetian and Late Devonian faunas from NW Westem Australia; and Early Devonian faunas from two small areas of the South Island of New Zealand, primarily from the Reefton area. All represent low latitude environments. Overviews of the biostratigraphy of eastern Australian rugose corals have been presented by Philip & Pedder (1967), Garratt & Wright (1988) and Zhen (1998). A small Frasnian fauna from Irian Jaya was described by Oliver ef al. (1995) who recognised aft1nities with the low diversity Frasnian faunas fi'om Western Australia.

Biogeographic patterns based on the distribution of mgose corals have traditionally been interpreted from a Northern Hemisphere perspective (e.g., Oliver, 1977, Oliver & Pedder, 1979; Pedder & Oliver, 1990). Australian rugose coral faunas are dominated by genera common in W Europe; it has been traditional to regard them as having Old World Realm affmities. In a recent compilation Zhen (1998) listed 97 genera of rugose corals documented from Australasian Devonian sequences and discussed their biogeographic implications. Zhen et al. (2000), drawing on this database, discussed evolutionary innovations, diversification patterns, turnover rates, decreases in diversity, and the effect of extinction events in E Australia. Biogeographic differentiation within the E Australian faunas is not obvious for any interval of Devonian time.

Of the 97 genera identified to date, 18 were based on Australian species. Many semicosmopolitan genera originated in eastern Australia in the Early Devonian, subsequently spreading to other regions, whereas Givetian faunas were marked by an ample influx of genera from southern China, central Asia, northwestern Canada and the Old World Reahn. Diversity was high in the Early Devonian-reaching a


maximum in the Emsian-and again in the Givetian, but was low in the Eifelian and Frasnian.

Lochkovian faunas are poorly known, although limestones of Lochkovian age are known from conodont data from eastern Australia: from the upper part of the Elmside FOlmation at Yass, the Camelford and GatTa Limestones at Wellington, Eurimbla, The Gap and Manildra, and from the Clandulla Limestone and Yellowmans Creek FOlTllation in the Cudgegong-Kandos districtall in east-central NSW. These faunas are essentially undescribed but include species of LYlielasma, Aphylilim and Rhizaphylilim.

Pragian faunas are known from numerous localities in eastern Australia. Characteristic genera are Carlinastraea, Paradisphyllum, Sterietazphylllllll and Martinaphylilim. In his survey, Zhen (1998) lumped late Lochkovian and Pragian faunas into a single undifferentiated unit consisting of 49 genera of which 40 appeared in this interval. Seven (possibly nine) genera are endemic to Australasia, though there are strong ties with Canada-13 out of 30 genera are common to the two regions. A biogeographic link between the South China and Tasman Regions is indicated by the shared occurrence of the operculate coral Chakeola (Wright, 2000) from the Emsian of Vietnam and from the late Lochkovian to Emsian ofNSW and Queensland. It probably includes all Pragian forms of 'Caleeala' repOlted from E Australia.

Emsian faunas, known from Victoria to nOlih Queensland, include 53 genera, many of them newly evolved forms. Phillipsastreids, stringophyllids and ptenophyllids are prominent. Distinctive endemic genera include FromeaphyllulII,Melrosia, Melasmaphylilim and Sanidaphyllum. Numerous genera (e.g., Mansuyphyllum) are shared with Europe and Southeast Asia. Several genera known from the Emsian of E Australia occur in younger strata elsewhere. These faunas display llukages to North America (25%) and South China (53%). An undescribed new genus of operculate coral, based on only three specimens, is known from the Emsian of the Taemas area and the Montagne Noire of southem France.

Faunas from the Reefton Group (probably Emsian) of New Zealand and the Cavan Bluff Limestone (earliest dehisee"s Zone) of the Taemas area, NSW, contain TipheaphyllulI1. Though the Reef ton faunas have semicosmopolitan genera of brachiopods and bivalves, they share several genera with the cold water Malvinokaffric Reahn (see other sections of this paper).

EifeHan rugose coral faunas-a mere 20 genera-are known from the Tamworth Belt of


NE NSW and NE Queensland. Genera common elsewhere do not appear in E Australia; important endemics are Sanidophyllutll, Blysmafophyllutll and possibly Amal'Ophyllum.

Givetian faunas, documented from the same regions, are more diverse and less provincial than Eifelian and older faunas, increasing to 44 genera with prominent immigrants from NorthAmerica, Europe, S China and Canada. Of the 36 genera described from the Fanning River Group ofNE Queensland (Zhen & Jell 1996), 77% occur also in S China, 72% in Europe and 39% in Canada. This is in marked contrast to the previous pattern of evolution of new genera in W Australia and their subsequent migration elsewhere. The Givetian faunas of West em Australia (15 species) are characterised byptenophyllids and disphyllids whereas the Frasnian faunas are dominated by disphyllids and phillipsastraeids. VelY few ofthe W.A. Givetian species are known outside the Cmming Basin. No species are known to extend from the Givetian into the Frasnian, doubtless a reflection of the Taghanic Event.

The only Frasnian fauna (12 genera) documented from E Australia is from the Mostyn Vale FOlmation ofNE NSW. It is less diverse at generic and specific level than coeval faunas from WestemAustralia (19 genera); 67% of its genera occur in WestelTI Australia as well (Brownlaw 2000). Frasnian faunas are more cosmopolitan than the pre-Frasnian faunas in Australia; Hillasfl'aea from Western Australia is one of a very few endemic genera. The Canning Basin faunas are generally oflower diversity than coeval faunas from S China or W Canada.

Famennian faunas are known almost exclusively from basinal deposits in Westem Australia. Very few platform-dwelling genera survived the Upper Kellwasser Event at the Frasnian-Famennian boundmy.

Ta bula ta ( G. Dargan) Tabulate corals are remarkably abundant and

diverse in the Early and Middle Devonian of Australasia but have been neglected taxonomically during the past 30 years compared with initiatives elsewhere, especially in the fonner Soviet Union, Vietnam, Mongolia and China. Theil' palaeobiogeographic potential remains largely untapped.

Leleshus (1970) concluded that there were four tabulate-based Ludlow palaeobiogeographic provinces, declining to a single worldwide province by the beginning of the Early Devonian. Dubatolov (1972), however, suggested that 12 provinces could be discdminated dming the Early Devonian; these he grouped into four entities corresponding approximately to the four Ludlow

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provinces recognised by Leleshus. Dubatolov's Australo-Eurasiatic group extended over E Australia and Asia, including the Asiatic part of the fOlmer USSR.

Occurrence of three Early Devonian tabulate genera in Australia accords with Dubatolov's analysis. Thecostegifes appeal~ in the Pridoli in Tadjikistan (Chekovich, 1960), and the Polar Urals (Chudinova, 1986); it enters at the base of the Pridoli inNSW (Dargan, 1993). It also occurs in the Point Hibbs Limestone (Pragian) in Tasmania (Jell & Hill, 1970) and in the Pragian in Vietnam and the Tien Shan, but did not appear in Europe and NorthAmerica until the Middle to Late Devonian, persisting through to the Early Carboniferous in Asia.

Yaclitiopora appears to be confined to the Early Devonian of Asiatic Russia (AI'khovik, 1986), SE Asia (Tong-Dzuy Thanh ef al., 1988) and Australia. It has been recorded from the Douglas Creek Limestone (early Emsian) in central Queensland (Jell & Hill, 1969), the Windellama Limestone (Lochkovian) of SE NSW (Mawson, 1973), the Bennys Tops Limestone in the Pigna Barney area ofthe TamwOlth Belt, NSW (Dongal, 1994), and the Milpose Volcanics near Bogan Gate (Dargan, 1993). These units range in age from Lochkovian to Pragian and possibly early Emsian.

Pseudoplasmopora occurs in the Late Silurian inAsia, Europe and Australia but appears to have become restricted to Asia and Australia in the Early Devonian. A similar genus, Pachyhelioplasrna, occurs in the Early Devonian and possillly also the Middle Devonian if Asia. It is possible that at least some oftheAustralian Early Devonian specimens identified as Pselldopiasl11opora gippslandica are actually Pachyhelioplasma. For instance, P. gippsiandica repOlted by Dongal (1994) from Pigna Barney NSW has smaller tabularia than is typical for P. gippslandica and, with its thickened walls, is more typical of Pachyhelioplasl11a.

Most of the Late Silurian tabulate holdovers had become extinct in Australasia by the beginning of the Middle Devonian. The clastic Baton FOlmation of the NW part of the South Island of New Zealand, has yielded few corals, apart from the tabulate PleurodicfYlll11 megasfomum McCoy (Shhley, 1938), a distinctive species of a semi-cosmopolitan genus. P. megasfol11l1l11 is widely distributed in Early Devonian clastic sequences in SE Australia. The West European Roemeripora is common in Emsian faunas, e.g., from Loyola, from the Lilydale, Cooper Creek, MUlTindal and Buchan Caves Limestones of Victoria, from the Douglas Creek Limestone of central Queensland, and from

200

the Garra and Jesse Limestones and Sutchers CreekFmmation of east-central NSW (Hill & Jell, 1970; Hill, 1978). Middle Devonian faunas of tabulates, for example from the Moore Creek Limestone of the Tamwotih Belt, are dominated by Favosites and Thaml1opora (BIiihl & Pohler, 1999) and had a distinctly cosmopolitan aspect. The beginning of the Late Devonian saw extinction of the Heliolitina and most of the Favositidae, Alveolitidae and Pachyporidae. ThalJ111opora-dominated tabulate faunas became an increasingly minor component of coral faunas.

Bryozoa (J. Phillips-Ross) There has been no previous synthesis of the

biogeography of Australasian Devonian bryozoans apari from a brief account presented by Bigey (1985) in her global analysis of the biogeography of Devonian bryozoans.

Bryozoans from the Devonian oftheAustralian part of Gondwana consist mainly of endemic species and are remarkably limited in occurrence. They are absent or sml'risingly rare in the large tracts of platform clastics and carbonates (e.g., the Buchan-Taemas-Molong Platfotm) in the Lachlan Fold Belt between Tasmania (e.g., Gill, 1950) and NE Queensland. Faunas repotied have generally few taxa, and often few individuals. An exception is a diverse early Emsian fauna of trepostomids, cyclostomids and ctenostomids described by Talent (1963) from the Wentworth Group in the watersheds of the Mitchell and Wentworth Rivers ofE Victoria.

Lochkovian-Pragian species fi'om the Lilydale and Loyola areas in Victoria include a cryptostomid Fenestella margaritifera and a cystoporate Fistulipora victoriae (Ross, 1961). A few bryozoans are known from the Pragian pari of the Garra Limestone of east-central NSW, notably in the Wellington and Emimbla areas, but these have not been described.

Early Emsian bryozoans have been known from several localities in the limestones of the Taemas-Wee Jasper area. For example, from near Taemas, Crockford (1941) described SemicosciniulJ1 vallatum from the Receptaculites Limestone (Ems ian) and, from nearby, a trepostomid CyphO/iypa(?) shearsbyi. From the same Taemas-Wee Jasper area Ross (1961) described the trepostomids Cyphotrypa mlll'rumbidgensis, C. lamellosa, Sterotoechlls shearsbyi, Homotrypa? sp., and Heterotrypa pontensis; the cryptostomids Ikelarchimedes warooensis, Hemifl)pa sp. A, Hemif1ypa sp. B and Nicklesopora geuriensis; and the cystoporate Fistulipora norensis. Typical of sparse OCCUlTences in the Lachlan Fold Belt is Pedder's (1971) report of a trepostome from the early


Emsian Lickhole Formation in the Ravine area of SE NSW. The only other bryozoans from this superbly exposed sequence appear to be undescribed species of Fenestella.

Blyozoans are a rare, low-diversity component of the large, often highly fossiliferous developments of carbonates in the Tamworth Belt of northem NSW, and in the Burdekin Basin, Broken River and Hodgkinson regions of NE Queensland. Only one fmID has been described from these regions: the cryptostomid Fenestella mOl/ara described by Crockford (1941) fium the Eifelian~arly Givetian Moore Creek Limestone notih of Tamworth.

In the Camring Basin, Western Australia, the cystoporate bryozoan Fistulipora pillarensis occurs in the later part of the Givetian (or possibly the early part of the Frasnian). Higher in the succession, a Frasnian bryozoan fauna comprises a cystoporate Fistulipora sadlerensis and the cryptostomids Fel1estella emanuelana, Nicklesopora westralis, N jitzroyensis and N leopoldensis.

The highest part of the Canning Basin succession, characterised by the Avonia proteus Zone (Famennian but not latest Famennian) has an abundant fauna of corals, nautiloids, brachiopods, and bryozoans. These bryozoans include a biogeographically distinct group of stenoporids: Percyopora tubulata, P. occidentalis, Fitzroyopora oscarensis and Granivallum jistulosum, along with the cystoporate Coe/ocaulis maculosa and the cryptostomids Nicklesopora crenulata and Fenestella pikerensis.

It is not sU1l'rising that bryozoans from E Australia might at first appear biogeographically distinct from faunas from Westem Australia. Bryozoan faunules described fium EAustralia are almost exclusively Emsian or older (only one species described from a post-Emsian horizon) whereas the Western Australian faunas are exclusively Late Devonian in age. Meaningful comparison of bryozoan faunas between E and W Australia requires a greatly enhanced database, especially with regard to the Emsian-Givetian of the Tamworth Belt of NE NSW, and the Lochkovian~arly Frasnian of the Broken River region and late Emsian-Famennian of the Burdekin Basin ofNE Queensland.

Bivalvia (M. Bradshaw) Although the presence of bivalves in the

Devonian sequences of Australia and New Zealand has been noted by many authors (Table 3), they are sparse in number, frequently poorly preserved and, in many cases, inadequately described and figured. Only recently have well preserved specimens becOlne available for

AAP Memoir 23 (2000) 201

FORMATION LOCALITY AUTHOR

Winduck Group White Cliffs Dun, 1898

Various Melbourne, Victoria Chapman, 1908

Reefton Group Reefton, New Zealand Allan, 1935

Baton Formation Baton River, New Zealand Shirley, 1938

Ruddocks Siltstone near Lilydale, Victoria Gill,1940

'Kimberley Limestone' Western Australia Teichert, 1943

Reefton Group Reefton, New Zealand FleminQ, 1957

Buchan Caves Limestone near Buchan, Victoria Talent, 1956a

Marble Creek Limestone near Walhalla Talent & Philip, 1956

Burdekin Limestone Queensland Heidecker, 1959

Coopers Ck Fm; Boola Beds Tvers reqion, Victoria Philip, 1962

Tabberabbera Formation Mitchell-Wentworth Rivers, V. Gill, 1949; Talent, 1963

Mcivor Ss & Mt Ida Fm Heathcote, Victoria Talent, 1964

Etonvale Formation Queensland McKellar 1966b

Adam Mudstone Reefton New Zealand Bradshaw 1974

Barrow Ranqe Beds Co bar NSW Fletcher 1975

Cavan FmlTaemas Ls/Buchan NSW and Victoria Johnston, 1993 Go Reefton Group Reefton, New Zealand Bradshaw, 1999

Table 3. Localities of salient Devonian bivalve faunas from Australia and New Zealand.

taxonomic and morphologic studies that can assist biogeographic analysis in a global context. The best Early Devonian bivalve fauna from SE Australia and the South Island of New Zealand are from within the Tasman Region, an entity originally inumed from brachiopod data.

The bivalve faunas from E Australia contain many endemic species and some endemic genera reflecting the well expressed Early Devonian provincialism, reaching an acme during early Emsian time. Conspicuously, the typical Northern Hemisphere palaeotaxodont Nuculoidea is replaced by Notonucula. The latter is well represented in Antarctic Emsian sediments (Bradshaw & McCartan, 1991) as well as in New Zealand (Bradshaw, 1999), and possibly in Victoria, Australia (pers. obs.). In limestone facies, however, Notollucula seems to be replaced by Nuculopsis (Johnston, 1993).

A rich fauna of silicified bivalves from Emsian limestones near Taemas, NSW, and The Basin, near Buchan, Victoria, was described by Johnston (1993). The 39 taxa described constitute the bestpreserved and generically the most diverse Devonian bivalve fauna yet found in Australasia. The palaeotaxodont component of the fauna comprises up to 60% of the bivalves fi'om some horizons. Johnston described 32 new taxa, many

of them belonging to typical Devonian bivalve genera, such as Cornellites, Actinopteria, Limopteria, Pseudoavicu/opecten, Goniophora, Mytilarca and Cypricardinia, all common to both the Old World andEastem Americas (Appohimchi) Realms, but much rarer in Malvinokaffric Reahn faunas. Johnston found Paracyclas proavia (Goldfuss) andP. rugosa (Goldfuss) in the Taemas Limestone to be indistingnishable from European and North American examples, with a North American influence reflected in the presence of Glyplodesma, G. buchanensis (Talent), in Victoria and NSW. Endemic genera include Nargunella Talent, together with a new and unnamed genus of Rhombopterii and an unnamed new genus of Lucinoidea.

A rich and well-preserved Emsian bivalve fauna fi'om clastics of the Reefton Group, New Zealand (Bradshaw, 1983, 1999) is dominated by palaeotaxodonts, with new spccies ofNatanucula, Nuculites, Ctenadontella, Paleoyoldia and Plteslia. Three species of Cornellites, one of Leptodesma (Leiopteria) and one ofGlyptodesma, all endemic to Reefton, reflect generic similarities with both the Old World and Eastern Americas (Appohimchi) Realms. Lyriopecten, represented by L. casterorulll (Fleming), is known elsewhere only in North America.

202

Strong affinities of the Reefton fauna to Victoria, already indicated by brachiopods, is sllppOlted by the presence of Nmgllnella ?nitida Talent in theAdam Mudstone at Reefton. A further link is shown by Gypricardinia crenislra (G. & F. Sandberger), an Old World Realm species, in both the early Emsian Tabberabbera Formation of Victoria (Talent, 1963) and the Bolitho Mudstone of Reefton. The presence of NlIclllodonta in New Zealand provides another indication of strong links with Europe, the genus being known elsewhere only in Sweden. NlIcllliles prirnus Bradshaw shows close similarities to NlIclIliles ellipliclIs of Europe.

The ReeftOll Group bivalve faunas are quite different from the Early Devonian bivalve fauna from the Baton FOlmation cropping out about 120 km fmihernOlth. Lochkovian conodonts from thin limestone lenses immediately below the base of the BatonFOlmation at some localities (J. Simes & R. Mawson, unpub. data) indicate that it is largely if not entirely Lochkovian in age. Recollection has shown that its bivalve fauna is prolific and varied. Compared with the Reefton faunas, Pseudoaviculopecten replaces LYl'iopecten and Pterinopecten, and the species of Gomelliles diller.

The relative positions of the Reefton and Baton centres of Devonian sedimentation are uncertain. Baton Formation faunas have strong E Australian and Old World affinities, with a possible close link with the Winduck Group of W NSW (Sherwin, 1990, 1995). The palaeotaxodont Glenodonla (Proeclenodonta), long thought to be endemic to Victoria (Philip, 1962), occurs also in the lower part ofthe Baton Formation (pel's. obs.). Although no connection is obvious between the Baton fauna and the faunas of the ?somewhat younger Reef ton Group, both regions can be argued to have had ease of communication with E Australia.

The presence of Malvinokaffric taxa in the Reef ton fauna (both brachiopods and bivalves) accords with this palt of New Zealand having been part of a discrete subprovince (Reeft011 Subprovince) lying much farther S of the Baton River area than at the present day, and isolated from it. Obrimia, a bivalve known only from Emsian horizons in Antarctica, has now been found at Reef ton; a form comparable to Modiomorpha herculi Bradshaw, a common Antarctic Devonian bivalve, is now known to occur at Reefton. Faunal links with Argentina along the Antarctic margin of Gondwana is suggested by the presence of NlIclIlites argentinum Sanchez in the Reefton Group, as well as the apparent presence of Glenodonlella gagei Bradshaw in Argentinian faunas. Palaeoneilo, a


common palaeotaxodont in South American and South African Early Devonian faunas, is absent from Reefton. The Reefton fauna contains two species of the endemic genus Paleodora (P. reeftonensis Fleming andP' angulata Bradshaw); this genus may be grouped at family level with Sil1odora, a South China genus.

The arenites of the Lochkovian Mount Ida Formation, particularly Unit 3 (Stoddart Member), of north-central Victoria contain a nearshore bivalve fauna first described by Talent (1964). The fauna, currently under re-study foIlowing re-coIlection, includes Gomelliles, Cypl'ical'dinia, ?NotonliGula, Phestia, Praeclenodonta, Goniophol'a and Cosl11ogoniophora.

Middle and Late Devonian bivalve faunas are not well known from Australia and are absent from New Zealand. A smaIl number of bivalve taxa occur in coarse, nearshore late Eifelian sediments in the Burdekin Basin of NE Queensland. These include the thick-sheIled Tanaodoll, formerly believed to be endemic to Australia; it is now known from species in the Middle Devonian of South China and Kazakhstan, one of which is identical with the Queensland form (Pojeta el al.,1985). Considering the cosmopolitan nature of Middle and Late Devonian faunas globally, the absence of Tanaodon in American and European faunas is surprising, although Johnston & Goodbody (1988) recorded a new Tanaodon-like bivalve from the Middle Devonian of Arctic Canada.

Teichert (1943) recorded a rich fauna of bivalves in late Middle? and Late Devonian limestones in the Kimberley region of Westem Australia. Pterioids are conspicuously absent; the bivalves present include Palaeoneilo, Goniophora, Mecynodon, Myalina, Macrodus, Schfzophoria, Buchiola, On/aria, Praecardium, Paracyclas and Eumega!odoll. The fauna has strong affinities with Europe and E North America; it includes bivalves characteristic of reef environments.

Rostroconchia It is interesting that the rastroconchs, referred

to broadly as COllocardiulJ'l, have numerous occunencesinEAustralia (Fletcher, 1943). They occur frequently but never in abundance in many silicified Early Devonian faunas from SE Australia but, curiously, have not been found in either of the New Zealand areas having Early Devonian sequences: Baton River and Reef ton.

Gastropoda (A. Cook) At species level, the Early Devonian gastropod

faunas ofE Australia (Chapman, 1916a; Talent,


1963, 1964; Talent & Philip, 1956; Tassell, 1976, 1977,1978,1982; Fan-ell, 1992; Cook, 1995) are strongly endemic although containing typically Old World Realm genera such as Mow'lonia, Bembexia, TrochoJlema, Euompha!opterus, Bellerophon, Tropidodiscus, Michelia and Naticopsis. Mitchellia was seemingly endemic to E Australia during the early Emsian.

The few documented Middle Devonian faunas (Etheridge, 1917b, 1921b; Heidecker, 1959; Cook, 1993, 1997; Cook&Camilleri, 1997) have a broadly cosmopolitan aspect showing some taxonomic linkages with European and North American faunas containing abundant palaeozygopleurids, gyronematines and murchisoniids. Three forms from the latest Emsian-earliest Frasnian Broken River Group of NE Queensland call for comment. The gyronematid Brokenriveria is also known fl.-om the Rhenish Givetian, a new genus of murchisoniid occurs in the Eifelian of Alaska (Blodgett & Cook, in prep.), and Denayella occurs also in the Eifelian ofN evada. The Late Devonian (Frasnian) fauna f\'om the Bonaparte Gulf Basin (Cook, 1998), includes Pseudomphalotrochus, Aglaoglypta and Plagiothyra, has strong links with European and North American faunas. Provincial or regional affinities are thus not clear.

Noteworthy undescribed faunas occur in the Lochkovian Windellama Limestone (Mawson, 1973; Mawson & Smith, in prep.), the late LOChkovian-Pragian part ofthe GalTa Limestone (Johnson, 1972; Mawson et al., in prep.) and the Brogans Creek Limestone, S of Mudgee, NSW, as well as the latest Lochkovian-early Pragian Martins Well Limestone Member of the Shield Creek FOlmation, NE Queensland. Undescribed Frasnian faunas from the Canning Basin include bellerophontids (e.g., Aglaoglypta), murchisoniids, suhulitids and microdomatids; these show affinity with faunas from the Iberger Kalk, Germany, the Voroneje Beds of the E Russian PlatfolTll, and the Naples fauna of New York. A Famennian fauna from the Canning Basin is under investigation (Cook, in prep.) and there are substantial undescribed faunas of Famennian age (marginifera and expansa zones) in the Burdekin Basin ofNE Queensland.

Palaeogeographic reconstructions (e.g., Scotese & McKerrow, 1990; Metcalfe, 1996) suggest greater ease of communication with major Asian blocks to the northwest of Australia and through them to Baltica (NW Europe) during Devonian times, but with a major gap across the HProto-Pacific" to Laurentia (North America). The presently available database for Australasian Devonian gastropods, though substantial, seems not to be sufficient for clear evaluation of the

203

significance of the "Proto-Pacific" gap in the biogeography of Australasian Devonian gastropods.

Non Ammonoid Cepbalopods (J.A. Talent) Non-ammonoid cephalopods are abundant in

some Devonian horizons in Australia, notably in: 1. Unspecified Early Devonian at Mt Matlock

in east-central Victoria: Plagiostomoceras thomas; Teichert & Glenister (1952).

2. The Buchan Group of E Victoria (Emsian, especially early Emsian deiliscens and perbonus zones; Teichert & Glenister, 1952): Buchanoceras graviventrum Teichert & Glenister, Litogyroceras spirale Teichert & Glenister and a remarkable suite of genera with actinosiphonate deposits: Pectinoceras subtl'igollum (McCoy), Polyelasmoceras adlillcum Teiche11 & Glenister, Brachydomoceras erectU1J1 Teichert & Glenister, and Macrodomoceras hOl-vitti Teichert & Glenister. Apparently similar faunas occur in the Taemas Limestone ofSE NSW but have yet to be investigated.

3. The Burdekin Basin (late Eifelian-Givetian) ofNE Queensland where the faunas have yet to be investigated.

4. The Late Devonian Gneudna Limestone of the Carnarvon Basin of Western Australia

5. The Late Devonian of the Canning Basin, WestemAustralia (Teichelt, 1939, 1940; Teichelt & Glenister, 1952): Michelinoceras cf. schlotheimi (Steininger), Galtoceras kirnbel'leyense Teichert, Wadeoceras australe Teichelt, COllstichoceras hardmanni (Etheridge) and CaYlitoceras iniqiseptatllm (Teichert)

The focus has been on previously undescribed genera; all described genera and species are endemics. Because of this, and because no data have been published in the past 50 years, the question of biogeographic pattems for this group cannot be addressed.

Trilobita (G. Edgecombe) Chatterton et al. (1979) reviewed the then

knownAustralianEarly Devonian trilobite faunas and reiterated the view of Campbell & Davoren (in Talent, 1972) that closest affinities are with "Old World" Early Devonian faunas of North Africa and central Europe. It should be noted, however, that this derives more from sharing a large number of genera-many of which occur also in other areas-rather than endemics. In most cases, since genera are widespread identification of sister species is required to determine relationships. Only a few Australian Devonian trilobites are endemic at the generic level.

Lochkovian trilobites are known from numerous units in SEAustralia. These include the

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lower, transitional part of the BeIl Shale in Tasmania (Gill, I 949b), the Stoddalt Member of the Mount Ida F onnation in the Heathcote district, (Holloway & Neil, 1982), the Humevale Siltstone near Lilydale (Gill, 1945a, 1949a; Shergold, 1968), the Melboume-Humevale transition beds, Christmas Hills (Jell & Holloway, 1983), and from allochthonous limestone in the Wurutwun Fonnation near Toongabbie (Holloway, 1991)all in Victoria; the Elmside Fonnation in the Yass district (Chatterton, 1971; Sherwin, 1971); and the lower part ofthe Tangerang Fonnation in the Bungonia~Windellama area, (Jones et al., 1986)-both in NSW, the latter arguably early Pragian. The cheirurid Azyptyx, from the Wurutwun Fonnation, is the only endemic genus. Most genera in the Mount Ida fauna are widely distributed (as are Zlichovaspis andReedops ii-Oln the Melboume-Humevale transition beds), but some informative species comparisons can be made, e.g., species of Scutellum and Crotalocephalus bear closest comparison to species from the early Lochkovian Kokbaytal Horizon in Kazakbstan (Holloway & Neil, 1982). The Australian OCCUlTence of acastids (Acaste and! or Acastella) is restricted to Lochkovian strata of Victoria and Tasmania, with species in the Bell Shale, Mount Ida Formation, and Humevale Siltstone. These acastids provide a useful stratigraphic and geographic link with closely allied taxa in the Baton FOlmation, New Zealand (Wright, 1990). Sthenarocalymene (=Apocalymene) from the Baton Formation (Shirley, 1938) is also most closely related to species from Victoria (S. angustioJ'; Humevale Siltstone) and NSW (S. australis; Elmside Formation).

A biogeographically infonnative element ofthe trilobite fauna ofthe Biddabirra Formation ofthe Amphitheatre Group near Cobar, W NSW, is a species of COI'dania, a genus otherwise endemic to the Lochkovian and Pragian of the Appalachian Province and Lower-Middle Devonian strata in Inner Mongolia and western Junggar, Xinjiang (Ebach & Edgecombe 1999). The status of the lichid Craspedarges in its type stratum, the Amphitheatre Group, is uncertain (Thomas & Holloway 1988). Taxa framJapan (Kobayashi & Hamada 1977) and the Qinling Mountains, China (Zhou 1987) referred to Craspedmges are now considered to represent a relict distribution ofthe mostly Silurian Borealmges Adrain, 1994.

Though a substantial part of the Garra Limestone of east-central NSW is late Lochkovian, silicified trilobite faunas from this unit at Wellington, NSW, are from Pragian horizons. Biogeographically restricted genera include the odontopleurid Laethoprusia, a


distinctive clade elsewhere known only from the Wenlock of Gotland and the Montagne Noire (Ramsk51d, 1991). A Pragian phacopid ft'om the Rosedale Shale neal' Limekilns, NSW (Wright & Haas, 1990) may be allied to other Australian Paciphacops.

Early Devonian falmas ft'omNSW and Victoria have closely related species in several faunas, for instance in Acanthopyge (Lobopyge), Kettneraspis, Dud/eyaspis (=Taemasaspis), Kainops, Paciphacops, Sthenarocalymene, Grota/acephalus, Liohal'pes, Scutellum, Coniproetlls, Proetlls (= Devolloproefus), and Cyphaspis.

Significant Emsian faunas include the Taemas Limestone of the Yass district, NSW (Chatterton, 1971), the Jesse Limestone at Limekilns, NSW (Wright & Chatterton, 1988), a presumably allochthonous limestone block at Flutation Hill, Mudgee, NSW (Chatterton & Wright, 1986), the Murrindal Limestone near Buchan, Victoria (Holloway, 1996), and the Tabberabbera Fomlation in the Mitchell-Wentworth rivers area, Victoria (Talent, 1963). Endemic genera are recognised within the Styginidae (Dentalosclitelllllll andA)'oeax), although each of these is monotypic. Wright & Chatterton (1988) obselved similarity between the Jesse Limestone fauna and those of Bohemia but cautioned that this may reflect facies control more than biogeographic affinity. Nevertheless, some genera in the fauna (e.g., AlberticOI),phe) are otherwise restricted to Bohemia and North Aft·iea. Some lichids in these Emsian faunas have informative dish'ibutions: Acanthopyge (Jaspel'ia) is resh'icted to Taemas and the Eifelian of Japan, whereas Terranovia from the Mudgee fauna is otherwise known only from N Canada and the Uralian Province (Novaya Zemlya, the Salair and the Altai and Omulev Mountains) in Lochkovian to Emsian horizons.

Detailed phylogenies that may be applied to biogeographic analysis are available for only a few groups represented in the Early Devonian, including phacopids (Ramsk51d & Werdelin, 1991) and odontopleurids (Ramsk51d & Chatterton, 1991). Australian LochkovianPragian Paciphacops fOlms a monophyletic group that is nested within and most closely related to Appalachian species. The only Malvinokaffric members of Paciphacops, from the basal Lochkovian in Bolivia and Argentina, ally with the North American, rather than Australian, species (Edgecombe & Ramsk5ld, 1994). Unlike a similar Appalachian/Australian pattern for COl-dania, the Australian lineage within Pacip/tacops had diverged by the Ludlow (it includes P. latigenalis, fram the Rainbow Hill


Marl near Yass, NSW). Australian Lochkovian Kainops species are also nested within an Appalachian Siluro-Devonian group, but are even more closely related to species from the Pragian of Bohemia. Leanaspis as redefined is endemic to the north margin of Gondwana (Armorica, Morocco, Turkey, Australia), the sole Australian representative beingL.jenkinsi fl·om the Elmside Formation (early Lochkovian, NSW). Several Australian Early Devonian species formerly assigned to Leonaspis are now referred to Kettneraspis. They do not appear to form a monophyletic group, with K. clavata from the Taemas Limestone being most closely allied to the Turkish Eifelian K. leucathea (Ramskiild & Chatterton, 1991). In summary, available c1adograms indicate Appalachian and North Gondwana affinities for different elements ofthe Early Devonian faunas.

The only Middle Devonian trilobites recorded from Australia are from the Broken River Group of N Queensland (Feist & Talent, 2000). Some taxa from the Broken River sequence are very closely related to forms from the Zhusileng Formation (Emsian) and Yikewusu Formation (Eifelian) of Inner Mongolia (Zhou et aI., 2000). Biogeographic affinities between the Broken River and Inner Mongolian trilobites are shown by late EifeHan species of Proetlls (Devanaproetus) and latest Emsian or earliest Eifelian Burgesina.

Late Devonian trilobites are all but unknown from E Australia-Wright (1988) recording an indeterminate phacopid from near Barraba, NSW}-but are well represented in the Canning Basin. Late Frasnian faunas from the Canning Basin of West em Australia, cUlTently under study by R. Feist and K. McNamara, include genera also known fl·om Europe, including Scutellum, Dudleyaspis, Harpes, 'Olarion', C1J'phops, Palpebralia, and Pteroparia. Two species of Palpebralia are regarded as identical with those from the Montagne Noire (R. Feist, pefS. comm.), the only instances of such widespread conspecificity known for Australian Devonian trilobites. Pteroparia and Palpebralia are primarily restricted to the mid-late Frasnian of North Gondwana-both occurring in the Harz Mountains, Montagne Noire, central Moroccan Meseta, but also in the Rhenish Slate Mountains, withPalpebralia occurring in the Carnic Alps (R. Feist, pers. comm.).

Early Famennian trilobites from the Virgin Hills Formation in the Canning Basin (Feist & Becker, 1997) include an endemic phacopid, Babinops, but species of Trimerocephalus and Cyrtosymbole are closely allied to European species.

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Ostracoda (P.J. Jones) Of all the known Palaeozoic ostracods in

Australia, Devonian species number the most (103), with a maximum (79) described fl·om the Early Devonian rocks in the SE part of the continent (Chapman, 1903, 1904, 1912b, 1914b, 1916b; Kriimmelbein, 1954; Pribyl, 1962; Talent, 1963, 1964; Willey, 1970; Siveter, 1976; Reynolds, 1978; Copeland, 1981). They are also described from Early Devonian rocks of the Georgina Basin (Jones in Turner ef 01., 1981), and Late Devonian sequences of the intracratonic basins (Bonaparte and Canning Basins) of WestemAustralia (Jones, 1962, 1968, 1974, 1985, 1987). Although many species remain undescribed, patiicularly in the Middle Devonian, the database is adequate to make some broad, qualitative, biogeographic interpretations for the early and late parts of the Devonian; these suppOli general observations on provinciality13.

In the Early Devonian, the earliest (Lochkovian) ostracod faunas are poorly known, except for the endemic BUllgollibeyrichia. This representative of the palaeocopid association ranges fi·om the top of the Bungonia Formation (about the Pridoli-Lochkovian boundary), near Goulburn, NSW, into the Humevale Siltstone (Lochkovian) and possibly the Kilgower Member of the Tabberabbera FOlmation (earliest Emsian) of Victoria (Talent, 1963; Copeland, 1981).

The Pragian ostracod fauna of the Lilydale Limestone, Victoria (Chapman, 1904; Willey, 1970) needs further taxonomic revision, but the described species (12) include representatives of the smooth podocopid association. At the species level, it appears to be endemic, the represented genera including Bairdia, Bairdiocypris, Microcheilille/la, Coelollella and other paraparchitaceans. The Lilydale ostracod fauna also includes some undescribed species of the palaeocopid association. These are also present in the Garra Formation (latest LochkovianPragian), near Wellington, and in subsurface Amphitheatre Group (early Pragian), near Ivanhoe, NSW (Jones & Willey, unpub. data). A key palaeocopid element is the endemic Renibeyrichia, originally described from the Taemas Group (early to middle Emsian) in NSW (Siveter, 1976; Reynolds, 1978). The palaeocopid association is also represented by Geisina victol'iana and Kloedenella waratahensis in the early Pragian Waratah Limestone ofWaratah Bay, Victoria (Kriimmelbein, 1954; Jones, 1974).

The earliest Emsian ostracod faunas from the Buchan Caves Limestone, E Victoria (Kriimmelbein, I 954)-no ostracods are repolied from the lowest, possible latest Pragian portion of this fonnation-and the Bell Point Limestone

206

ofWaratah Bay, Victoria (Kriimmelbein, 1954), provide an example of inter-tenane comparison of species for the Molong-Monaro and Melbourne terranes respectively. Both faunas are ofthe palaeocopid association. The Buchan Caves Limestone fauna contains the endemic genus Chapmanites (3 species), and 9 endemic species of genera 'Aparchites' (2), Cavellina (3), Coeloenellina (1), Kirkbyina (I), Kloedenella (1) and Slilcella (1). The Bell Point Limestone fauna contains 9 endemic species of genera ElIkloedenella (5), Chapmanites (1), Dibolbina (1), Bairdia (1), and Renibeyrichia (1 ~ Masligobolbina sodaUs Kriimmelbein, 1954). Of a total of 23 taxa from the Buchan Caves Limestone (14) and the Bell Point Limestone (9), only one (Chapmani/es lophotlls) is common to both rock units.

From the early Emsian Receptaculites Limestone, Taemas, NSW, Reynolds (1978) recognised the most diverse of Australian Early Devonian ostracod faunas (33 species). She referred the fauna to two assemblages: a lower brachiopod/gastropod assemblage dominated by palaeocopids (Renibeyrichia, Rishona and eukloedenellids) and an upper bryozoan/sponge assemblage dominated by Coniferina. The last genus is endemic in a broad provincial sense, as are the genera AcinacibolbinG, Cavanites, Gyrgathella, and Batalaria; these also form patt of the blyozoan/sponge assemblage. Ostracods of the brachiopod/gastropod assemblage are similar to those of the palaeocopid association in the Bell Point Limestone, whereas those ofthe b,yozoan/ sponge assemblage, clearly a mixture of Wang's (1988) spinose-podocopid, and smoothpodocopid assemblages, are unlike any known Devonian assemblage from elsewhere in Australia. The blyozoan/sponge assemblage of ostracods, however, closely resembles the fauna ofthe Thmingian Ecotype (Becker, 1976) present in the uppelmost (late Emsian) part of the Fukuji Formation of central Japan (Kuwano, 1987). Species in common that indicate close provincial links are Acanthoscapha brevicristata Reynolds (~ Acanthoscapha spp. A, B of Kuwano), Acinacibolbina anteropinnata Reynolds (~ Schohariella sp. A of Kuwano), and Cavoni/es robustihamatus Reynolds (~Cavanites? sp. B of Kuwano). Other common species, e.g., Berounella spinosa, and Tricornina robusticerata, suggest more distant provincial links with Europe ('Baltic-British' Province), via the Cordilleran Province of the Old World Realm of Boucot (1975). Earlier zoogeographic links with the CardilleranProvince are indicated by the presence of Gyrgathella (2 species) and COlliferilla (1 species) in the latest Silurian (Pridoli) portion of


the Roberts Mountains Fonnation, central Nevada (Stone & Berdan, 1984). In addition, Yukonibolbilla, originally described from the Ludlow and Pridoli ofthe District of Mackenzie (Copeland, 1977), and later from the Pridoli and Pragian of central Nevada (Stone & Berdan, 1984; Berdan, 1986), is also present in the late Wenlock of mid-western NSW (J. Banos, 1990, unpub. data).

There is alsa a zoogeographic link, albeit weaker, between the Thuringian Ecotype of the late Emsian part of the Polzheluo Formation, Guangxi, South China (Wang, 1989), and that of the Receptaculites Limestone (Wang & Jones, 1993). The species described by Wang (1989) as Thuringobolbilla? crassa and Hanaites sp. respectively, are probably conspecific with Thuringobolbina australis and Cavanites robustihamatus described by Reynolds (1978). Another biogeographic link, at the species level, is between the late Emsian ostracods of Guangxi, and those ofthe Craven Peak Beds ofthe Georgina Basin. This is suggested by the presence of a taxon first questionably referred to Healdiallella iJlcollstans Polenova (Jones, in Turner et al., 1981), but now more appropriately referred to H. subdistillcta Wang, 1983.

The Cravens Peak Beds also contain the eridostracan Oyptophyllus (?Branchiopoda), a fmID widely distributed in Emsian rocks in South China (Wei, 1988),AImorica (Becker & Sanchez de Posada, 1977, Weyant, 1980), the Cordilleran province afthe Nmth American Plate (Copeland, 1977), and the nmthemmargin of Gondwana (Le Fevre, 1963). Like benthic ostracods, this fmm presumably had no pelagic larval stage, and could not have crossed deep oceanic barriers. Instead, they could have migrated during transgressive pulses along juxtaposed shallow shelves of adjacent blocks.

Less well known are the Middle Devonian ostracods of Australia, partly represented by an undescribed fauna straddling the Middle-Late Devonian boundary in the Gneudna Formation ofthe CamarvonBasin (Jones, 1974). The fauna includes Svantovites, indicating a link with W Europe (Armorica), and the eridostracan Oyptophyllus, present in coeval N orthAmerican and European faunas (Jones, 1962).

Published research on Australian Late Devonian ostracods has been concentrated on assemblages fi'om Western Australia, where 24 species, eridostracans excluded (Jones, 1962, 1968), have been described from the Bonaparte Basin (Jones, 1968, 1985). Some ofthese species have been recognised in the onshore Canning Basin (Jones, 1987), but many more (in the collections of the Australian Geological Survey


Organisation, Canberra) await description from both basins (Jones, 1974). The ostracods listed by Jones (1974) from the early Frasnian Sadler Limestone of the Canning Basin belong to cosmopolitan genera (e.g., Amphissites and Bairdia). A fauna from the late Frasnian Westwood Member of the Cockatoo Formation of the Bonaparte Basin contains species of Amphissites, Bairdia and iJldivisia either conspecific with, or closely related to, forms recorded from the Frasnian of the Russian Platform (Jones, 1968).

The best known of the Late Devonian ostracod faunas of West em Australia is one present in the latest Famennian ('Stmnian') Buttons Beds, in the Bonaparte Basin (Jones, 1968). Although some taxa in this fauna have been revised (Jones, 1985, 1987), more revision is necessary; more species await description. The Buttons Beds fauna is highly diversified; it includes cosmopolitan genera of the palaeocopid and smooth podocopid associations (e.g., Chamishaella, Shishaella, Cavellina, Sulcella, KnoxUes, Mwginia, illdlvisia, Bairdia and Bairdioeypris). Many of these taxa are either conspecific with, or closely related to, fOlms recorded from Europe, Russia, Kazakhstan, the Kuznetsk Basin, South China, and North America. Genera that at present appear to be endemic are Notoscapha and Parabollchekius (Jones, 1985). Of interest is the presence of Kalalona in the late Famennian (,Stmnian') of the Bonapatte Basin and Belgium, indicating a zoogeographic link with Armorica (Jones, 1985). The ostracods, lacking a pelagic larval stage, probably indicate that the shallow shelves ofthe western part of Gondwana and Laurentia-Baltica were close enough to pennit genetic exchange.

In summary, the provincialism of the Early Devonian ostracod faunas of SE Australia is indicated by the endemic genera Blingonibeyri eh i a (Lochkovian-Pragian), Reuibeyrichia, Acinacibolbina, Cavanites, Chapmanites, Gyrgathella, Coni/erina and Bamlaria (Emsian). These are diagnostic ostracod genera for the Tasman Region, some of which demonstrate zoogeographic links with central Japan, South China, and the Cordilleran Province of the Old World Realm (Boucot et al., 1967; Boucot, 1975).As the benthic ostracods could not have crossed deep oceanic baniers, some genetic interaction between these regions is indicated, conceivably by migration along juxtaposed shallow shelves of adjacent blocks during transgressive pulses. The observed zoogeographic links may reflect existence of 'staging points' (isolated terranes) in the Panthallassan Ocean. The deat1h of Middle Devonian ostracods inAustralia does not allow any conclusion to be drawn about

207

biogeography of that time. By the latest Devonian, howevC1; there is sufficient evidence from benthic ostracods to suggest that they could have migrated along juxtaposed shallow shelves of the adjacent blocks of Gondwana and Laurentia-Baltica. A similar connection probably existed between the South China Plate and NW Australia within the eastem end of the palaeo tethyan equatorial belt.

Echinodermata (J.A. Talent) Devonian crinoids have been described from

12 areas inAustralia (Jell, 1999a; Jell & Jell, 1999; Webster; 2000):

I. From the Humevale Siltstone of central Victoria from several localities scattered from Lilydale westwards (Jell & Jell, 1999)

2. From the Mt Ida Formation (Lochkovian; apparently from unit 3-Stoddart Member-of the formation; Jell, 1982, 1999): Crotaloerinites pulcher (Hisinger) and Ctenoerinus pauGidaetylus Hall.

3. From two limestone olistoliths, the old Toongabbie Marble Quarries (probably early Pragian; Mawson & Talent, 1993), in east-central Victoria where calices arc relatively common. Three species have been described (Philip, 1961): Euealyptocrinites inehoatlls Philip, Thylacocrinus? ignotlls Philip and Hexacl'inites? sp.

4. From the upper part (early Pragian, sulcatus Zone) of the Martins Well Limestone Member of the Shield Creek FOlmation of the Broken River area of NE Queensland: Pandanocril1l1s martinslvellensis Jell et al. and Parapisocrinlls sp. (Jell et al., 1988; Jell, 1999b).

5. From the Emsian (probably kindlei Zone) Lilydale Limestone of central Victoria (Bates, 1972): PernerOCrill11S discus. This bizane genus, being known otherwise only from Bohemia (Bouska, 1947), is a good link to the BalTandian Old World Realm faunas.

7. From a limestone olistolith of assumed late Pragian age from Loyola (Mawson et al., 1992), near Mansfield Victoria: Eucalyptocrinites Jonzi Jell el al.

8. From high in the GalTa Limestone from a transgressive (deepening) horizon at Wellington, NSW, believed to be probably early Emsian dehiscens Zone because of occurrence of Polygnathus cf. P. pireneae from considerably lower in the Garra Limestone at "Mountain View" (R. Mawson, unpub. data). There are six named forms from this interval (Jell et al., 1988): Eucalyptocrinites rosaceOliS Goldfuss, Atielocrinites salus Jell et al., Pandanocrinus wellingtonensis Jell el aI., P. gellriel1sis Jell et al., ShimantocriJlus distinctodorslis Jell et al. and Strllszicl'inliS dulciculus Jell et al. Five ofthe six species and the last two genera were regarded as

208 AAP Memoir 23 (2000)

Fig. 13. Some endemic placodenns of East Gondwana (from Long, 1993, Fig. 9.6).


new and, having not been reported from elsewhere, are thought to be endemics.

9. From the 'Mount Holly Beds' (limestone olistoliths of Emsian age in the Late DevonianEarly Carboniferous Mt Alma Formation; P. Blake, pers. comm.) of the Rockhampton area of central E coast Queensland: Pandanocrinlls cf. martinswellensis Jell et al. extends the range of this genus, seemingly endemic to E Australia, from early Pragian to late in the Emsian.

10. From the Papilio Formation (Givetian; Mawson & Talent, 1988; Sloan el al., 1995) of the Broken River area ofNE Queensland. Of the four described species (Jell el al.,1988)Cllpressocriniles abbreviailis (Goldfuss), Rhipidoc17'l1US crenatus Goldfuss, Doia!oClinlis peregrill11S Jell et al. and Melocrinites tempestus Jell el al.-two were previously known from Germany. This accords with a general low provinciality of shelly faunas as a whole during Givetian times.

11. From the Late Devonian reef complexes along the nOlthern margin of the Canning Basin (Jell & Jell, 1999)

12. From the Campwyn Formation (?Late Devonian) near Proserpine, NE Queensland, a report of Me locri niles lempesilis Jell el al. (Jell, 1999c).

Only one crinoid has been described from New Zealand (Prokop, 1970), Megislocrinlls reeftonensis fi:om the Reefton Group. It has little biogeographic significance; Megistocrinlls is a long-ranging semi-cosmopolitan genus. Crinoid calices are known from other areas including the Late Devonian of the Canning Basin (Jell & Jell, 1999) and the Bonaparte Gulf Basin but faunas from the latter have yet to be described (p. A. Jell, pers. comm.). Disarticulated crinoid remains are often abundant, taxonomically diverse-almost 100 Early and Middle Devonian parataxa are known from E Australia-and display closest relationships with faunas described Ii'om central Asia (SlUkalina & Talent, unpub. data). This could well be a 'monographic effect' due to a major focus on disarticulated crinoid remains by workers in the fonner Soviet Union.

Attention is drawn to the remarkable mitrate homolozoan family Allanicytidiidae first reported-Allanicytidillmjlemingi Caster & Gill (l967}--from the Reefton Group (?Alexander Mudstone; M. A. Bradshawpers. comm.) in Rainy Creek, Reefton, New Zealand. A trans-Tasman linkage has been revealed by occurrence of the family in the Silurian of Tasmania and Victoria (Ruta & Jell, 1999a, 1999d). The Early Devonian (Lochkovian) Humevale Siltstone at Kinglake and Kinglake West, Victoria, has produced a remarkable diversity of carpoids, starfish and

209

ophiuroids. From it the homolozoan echinoderms Rlilrocopelis junori (Withers), R. vicloriae Gill & Caster, 'R.' wilhersi Gill & Caster, Adokeloempus janeae Ruta & Jell, Vicloriacystis ho/mesoJ'u1tl Ruta & Jell andPseudovictoriacystis problemaliea Ruta & Jell have been described (Gill & Castel; 1960; Jell & Holloway, 1982; Ruta & Jell, 1999b, 1999c). The above genera of homolozoans appear to be endemic to the Tasman Region. Starfish, earlier studied by Withers & Keble (1934a, 1934b) from the same area are in need of revision.

A major contribution to knowledge of the Lochkovian and ?Pragian crinoids of central Victoria has been presented by Jell (1999). Twenty one taxa identified to species-level have been reported, principally from various horizons of the Humevale Siltstone and from unit 3 (= Stoddart Member, Edwards el al., 1998) of the Mt Ida Formation. All were described as new except for Clenoerinus palleidaelyilis (Hall) and Phimocrinlls amer/canus Springer, forms with eastern NorthAmerican affinities, and C. stellifer Follman and C. signaills Schmidt, fonns with west European affinities. Curiously, among the 11 genera of crinoids described li'01n the Bokkeveld Group ofSouthAliica (Jell & Theron, 1999), only one genus, OphiocJ'inlls, occurs as well in the Early Devonian faunas from central VictOlla. This contrastrefiects Old World Realm biogeographic affinities of the central Victorian faunas and Malvinokaffric Realm affinities of Bokkeveld faunas, but may be excentuated by the substantial difference in age between the the faunas from the two regions: basically Emsian for the latter versus Lochkovian for the fOlmer.

In contrast with the crinoids, only a single echinoid species is known from the Devonian of Australia, Cavanechinlls ·warreni from the earliest Emsian Cavan Bluff Limestone Member of the Cavan Formation at Taemas, NSW (Brown, 1967).

Eleven genera of crinoids, together with a blastoid and a cyclocrinoid have been described from the Late Devonian reef complexes of the Canning Basin (Jell & Jell, 1999). The blastoid, Hyperoblasills buglensis Jell & Jell, is representative of a western North American genus. The crinoids are a mix of two endemic genera (Wacrinlls and Playfordicrinlls), and rather widely distributed genera which, as a consequence of addition of the recently documented Canning Basin occurrences, may now be constrmed as having been at least semi-cosmopolitan.

Vertebrata (G.C. Young, J. Long & C. Bunow) Devonian fish assemblages of the Australasian

region l4 are dominated by diverse and widely

210

-1cm


Fig. 14. Some endemic acanthodians, osteichthyans and chondrichthyans afEast Gondwana (from Long, 1993, Fig. 9.8).

distributed placodem)s or armoured fishes (Fig. 13), but with some significant remains of other major groups (Fig. 14)-the osteichthyans (bony fishes), early chondrichthyans (cartilaginous shark -like fishes), and acanthodians (spiny-finned fishes). Jawless fishes (agnathans), which in the Northem Hemisphere have an extensive fossil record dominated by several highly diverse armoured groups, are almost unknown from the

Devonian of Australasia, a major differencewhich calls for biogeographic explanation (see below). The majority of Devonian agnathan remains from East Gondwana are referable to one group, the thelodonts, preserved as isolated scales, and abundant in some microvertebrate assemblages (Tumer, 1982-97).

Devonian vertebrate localities in Australia are predominantly marine in the west (Camarvon,


Canning, and BonapaIie Gulf Basins; see Fig. 3), predominantly non-marine in the centre (Amadeus, Georgina, and Officer Basins); and both non-marine and marine in E Australia (Lachlan Foldbelt, Tasman Orogenic Zone; see Figs. 2, 4). A recent overview of 36 Australian localities is given by Young & Tumer (2000, fig. 1), with more detail for Western Australia, Queensland and southeastern Australia provided respectively by Long & Trinajstic (2000), Tumer et al. (2000) and Basden et al. (2000). Devonian vertebrates have also been reported from Irian Jaya (Turner et al., 1995) and New Zealand (Macadie, 1985, 1998). Included in the Australasianregion is the significant Middle-Late Devonian Aztec fish fauna from S Victoria Land, Antarctica (e.g., Young, 1989b, 1991a); it belonged to the eastern pOliion of the Gondwana supercontinent during the Palaeozoic.

Habitat al/{I dispel"sal of Devolliall fishes. Freshwater fish distributions provide crucial evidence for biogeographic studies of continental faunas (e.g., Myers, 1949; Darlington, 1957; Berra, 1981), but recognising 'primary division' fishes (entirely restricted to freshwater) in the fossil record is difficult when marine invertebrate fossils are absent. The lack of definitive criteria for distinguishing marine from non-marine depositional environments has engendered much discussion in relation to reconstruction of past geography (e.g., Janvier, 1985; Young, 1987a, 1990a; Van Der Voo, 1988). Some classic fossil fish assemblages of 'Old Red Sandstone' type in Europe (e.g., Goujet, 1984; Schultze & Cloutier, 1996), and also in Australia (young, 1999; Jones et 01., 2000), may have been subject to marine influence; analysis of empirical data for some well-known fossil fish occunences (e.g., Schmitz et al., 1991, Prichonnet et al., 1996) has not produced unequivocal evidence of a strictly freshwater habitat.

Thus, Devonian fish assemblages of Australia come from both obviously marine, and uncertain (marginal or non-marine) sedimentary strata, the fonner often limestones with abundant marine inveliebrates, the latter typically thick sandstone! siltstone sequences traditionally interpreted either as fluvial deposits, e.g., the Hervey Group of central NSW (Johanson, 1998; Young, 1999, and references therein), or as lacustrine deposits in a volcanic setting, e.g., the Mt Howitt province of east-central Victoria (Long, 1999, and references therein).

As discussed elsewhere (Young, 1981, 1987a, 1987b, 1989a, 1990a, 1995a; Rich & Young, 1996; Young & Janvier, 1999), the distribution of Devonian fishes is clearly non-random in

211

relation to palaeogeographic reconstructions. Controlling fuctors can be related to patterns of past geography (see below). Chance dispersal across oceans is excluded as a general explanation. The distribution pattem of individual taxa may provide important clues to their habitat and dispersal capabilities.

One ecological grouping includes fishes that lived in shallow marine environments, but had the capacity to cross marine baniers, as indicated by their widespread distribution. Although the Silurian-Devonian vertebrate faunas of continental Asia and East Gondwana are very different (young, 1995a, Young & Janvier, 1999), they include examples of this type - the closely related arthrodires Kweichowlepis (South China) and Buchanosfells (Fig. l3L), or the lungfishes Sorbiforhynchlls(Guangxi; Wang ef ai., 1993) and Pillararhynchlls and Dipnorhynchus from East Gondwana (dipnorhynchids have also long been known from Europe).

A second ecological grouping includes forros commonly preserved in non-marine deposits, but with widespread distribution also indicating a capacity for marine dispersal. Groenlandaspid arthrodires (Fig. l3C-E) are a good example (Ritchie, 1975; Long, 1995a). The antiarch Bothrioiepis is the most widespread Devonian fish, represented worldwide by over 70 named species (Young, 1988a; Johanson, 1998; Johanson & Young, 1999). Such forms clearly could cross marine barriers, although subgeneric pattems may be biogeographically significant, for example the species group of Bothriolepis charactedsed by the shape of the preorbital recess (Fig. 15E-F), so far known only from South China and East Gondwana. More difficult to assess are the only known armoured agnathans from Australia: Pfturiaspis (Fig. 15A) possibly endemic at class level (young, 199Ib), and the associated Neeyambaspis (poorly known from a single incomplete sandstone impression). The latter shows superficial resemblance to some of the galeaspid agnathans of South China (Fig. 15B-D).

The galeaspids of Asia exemplify a third ecological grouping of major taxa, with distribution limited to one or a few blocks or terranes, indicating inability to cross marine baniers-as is the case for some major freshwater fish groups today. Other examples are the armoured agnathans (osteostracans and heterostracalls) of Euramerica (unknown in the Siluro-Devonian of Gondwana), and the sinolepid antiarchs (Fig. 15G-J), known only from the Early-Late Devonian of E Asia, and the latest Devonian ofEAustralia (Ritchie et al., 1992).

Devonian fishes also contribute to the evidence of palaeolatitude for the Australasian region.

212 AAP Memoir 23 (2000)

Fig. 15. Various Devonian vertebrate taxa relevant to the question of Gondwana-Asia connections. Similar but distantly related armoured agnathans ofE Gondwana and Asia. A, endemic agnathan PUuriaspis doylei (dorsal reconstmction) B, headshield of Neeyambaspis enigmatica (a-b from Georgina Basin, \VQueensland) C-D, headshields of galeaspids Asiaspis expal1sa, and Sinoszec/w(Jllaspis yanmenpaensis from South China (A-8 modified from Young, 1991, Fig. 7; C, from Pan ef ai., 1975, Fig. 5; D, from Pan, 1992, Fig. 29). E-F, skulls of antiarchBothriolepis with similar prcorbital recess; B. s/raokuanensis, Middle Devonian, China (E, after Chang, 1963; Lill, 1973); B. karawaka, Middle-Late Devonian, Antarctica (F, after Young, 1988a). G-J, sinolepid antiarchs from S China and E Australia, readily characterised by unique ventral fenestra in trunk annour (from Ritchie et al., 1992); G, Xichonolepis qujingensis, Middle Devonian, China, ventral view; H-J, Gren/ellaspis branagani, Late Devonian, E Australia, ventral and dorsal views (after Ritchie et al., 1992; Young & Janvier, 1999).


Many assemblages come from limestones with carbonate bnild-ups in the Early and Middle Devonian of E Australia (e.g., Taemas, Buchan and Broken River regions; Fig. 2) and reef complexes in the Late Devonian of the Canning Basin, Western Australia (Fig. 3). These suggest tropical-subtropical climates from which a low palaeolatitude may be infened (probably less than the 40" S on palaeomagnetic maps; Fig. 16).

ModernAustralia is well known for its highly diverse birds, reptiles and marsupials, but its freshwater fish fauna is depauperate-due to aridity of the continent--compared to the high rainfall areas of SE Asia, Africa and South America. Young (1990a, p. 247) suggested a Devonian placement of Australia in the subtropical dry belt to provide a similar explanation for its impoverished pre-Emsian Devonian fish faunas. New discoveries of microvcliebrate remains suggest that this may apply only to the Silurian, where our negligible fossil fish record contrasts with the highly endemic and diverse fish fauna fi'om marginal and non-marine Silurian-Devonian deposits of South China, Europe and NorthAmerica. An equatorial position for the South China block would have provided sufficient rainfall to support one of the most diverse and endemic faunas known fi'om the entire fossil veliebrate record (young, 1990a).

Early Devonian /tllmas. Early Devonian faunas of our region are typified by the Wuttagoollaspis fauna in assumed non-marine (fluviatile) sequences W of Cobar, NSW (Ritchie, 1969, 1973; see Fig. 2), and in the Georgina (Dulcie and Toko Ranges) and Amadeus Basins of central Australia (Turner et al., 1981; Young, 1988b, 1991b; Fig. 3).

Siluro-Devonian microveliebrate assemblages from SE and NE Australia (Basden el al., 2000; Tnmer et al., 2000) include various form-taxa based on scales which are widely distributed elsewhere, bnt with some indication of South China connections. The actinopterygian Ligulalepis from the Murrumbidgee Group (Schultze, 1968), is also known from China (Bunow, 1994). Acanthodian scales resembling Poracanthodes qUjingensis from China have been identified in late Silurian limestones in NE Victoria and central-westem NSW (parkes,1995 and in Basden et al., 2000), and in the Silverband Formation (Lochkovian-Pragian) of the Grampians, W Victoria (BUlTOW, 1997a; Turner, 1986). Colquhoun (1995) reported abundant fish plates (not documented) from the Clandulla Limestone and overlying Yellowman's Creek Formation (elll'ekaensis-deita Zones) SE of Mudgee. Parkes (1995, and inBasden et al., 2000)

213

reported microvertebrate faunas from the Camelford Limestone (?lVoschmidti-delta Zones of east-central NSW and from Windellama S of Goulburn (ellrekaensis-delta Zones). Late Lochkovian-early Pragian faunas from E Australian carbonates include the Garra Limestone (parkes & Hocking, in Basden et al., 2000), Pt Hibbs Formation of SW Tasmania (BmTow et al., 1998), ConnemarraFormationSE of Mud gee (Burrow, 1996, 1997b, in Basden et al., 2000), and Mariins Well Limestone Member of the Shield Creek Formation, NE Queensland (Tmner et al., 2000). Based on accompanying conodont and invertebrate faunas, and the vertebrate taxa represented, these were nearshore environments. The Martins Well Limestone Member has been interpreted as a lagoonal deposit (Withnall et aI., 1993) but it and other limestone members of the Shield Creek Formation have extraordinarily high diversity invertebrate faunas consistent with nearshore situations, with "normal" circulation (Talent et al., in prep.). The vertebrate fauna includes a thelodont, several climatiid and ischnacanthid acanthodians, and four or more placoderms (Turner et al., 2000). The ConnemalTa Formation in central NSW was deposited more distally than the underlying Deniwong Group, but not in deep water (Shenvin, 1996). Its vertebrates show more diversity at high taxonomic level than the Martins Well Limestone Member, with the thelodont TIIl'illia sp. cf. T. australiensis, four acanthodian taxa Nostolepis guangxiensis, N. sp. cf. N. striata, Trundlelepis cervicostulata, Machaeracanthus sp., six or more placodenns, an early actinopterygian Terenolepis turnerae, an onychodont, a dipnomorph, and a putative osteichthyan Lophosteus? incrementus (Burrow, 1995a, 1995b, 1995c, 1996, 1997b; BUlTOW & Turner, 1998). The Garra Limestone has a less diverse vertebrate fauna, with perhaps two acanthodians, four placoderms, and similar osteichthyans to the Connemarra Formation (Hocking & Parkes, in Basden et al., 2000). The fauna of the Point Hibbs Formation of SW Tasmania is impoverished, with three or four acanthodian taxa, perhaps three placoderms and an onychodont (BmTOw el al., 1998). Lack of thelodonts, and the composition of the accompanying conodont and inveliebrate faunas, suggest that the latter two units were deeper water deposits than the Connemarra Formation or Martins Well Limestone Member.

Mid Pragian carbonates with conodont and microvertebrate faunas are velY rare inAustralia. The only indubitable kindlei Zone veliebrates are from the Coopers Creek Limestone ofthe TyersBoola area ofE Victoria (Philip, 1965; Basden, 1999b). Mawsonel al. (1988; Mawson & Talent,

214

400 Ma Early Devonian

365 Ma Late Devonian

1994a) suggested the lack of conodont-producing carbonates from the post-killdlei limestones of that area could have been connected with a regressive interval in E Australia. Latest Pragian/early Emsian (pireneae-dehiscens Zone) conodont and microvertebrate-bearing carbonates are widespread in SE Australia, and include the Buchan Group ofE Victoria, the Mumlmbidgee and Byron Range Groups of SE NSW (e.g., Basden, 1999a; Lindley, 2000), and several units in central NSW. Among the larter, limestones of the Gleninga Formation have produced a rich microveltebrate fauna with abundant scales ofthe thelodont T. australiellsis, the climatiid acanthodian Nostolepoides platymarginata, an ischnacanthid, an acanthodian, G0111phonchus?

A

B


Fig. 16. Distributions of biogeographically important Devonian fish groups on global reconstructions. A, 400 Ma (Early Devonian); B, 365 Ma (Late Devonian). Squares: ostcostracans (Euramerican province; Tuva); triangles: amphiaspids (Angaran block); stars: Asian endemics; open circles: West/South Gondwana (Malvinokaffric)~ closed circles: East Gondwana province (phyllolepid occurrences in B including Euramerica).

bogongensis, the actinopterygian Ligulalepis toombsi, and rarer taxa including perhaps three placodelms, an onychodont, a dipnomOlph, and the chondrichthyan Ohioiepis sp. Samples from the Troffs FOlTIlation produced a similar fauna.

Interpretingthese microvertebrate data in telms of regional communities, it appears that the level of species-diversity is comparable in both older and younger strata from similar environments, despite changes in faunal composition. Diversity seems greatest in slightly offshore assemblages, decreasing with water depth to lowest diversity in near-shore/inteltidal deposits.

Macrovertebrates from the Early Devonian marine limestones of SE Australia also include endemics, such as the acanthothoracid placo-


derms Brindabellaspis, Murrindalaspis and Weejasperaspis (e.g., Young, 1980; Long & Young, 1988), with more widespread fmms, such as the lungfishes (Campbell & Barwick, 2000, and references therein) and buchanosteid arthrodires (White & Toombs, 1972; White, 1978; Yonng, 1979; Long, 1984). As well as the Chinese Kweicholepis (see above), buchanosteids are recorded from the northem Gondwana margin (e.g., Jauf Formation, Saudi Arabia; KhushYeilagh Fmmation, han).

Middle Devonian fallnas. Marine fish faunas of this age are poorly known in Australasia, but include both endemics and widespread fmills. The Broken River Group of NE Queensland has produced the seemingly endemic antiarchs WlIl'lli/gulepis and Nawagiaspis (Young, 1990b) as well as large atihrodires known from elsewhere such as At/anlidoslells and Aspidichlhys (Tumer el al., 2000).

The Hatchery Creek fauna from near Wee Jasper (Young & Gorter, 1981), of assumed EifeHan age, is a non-marine assemblage including widely distributed taxa such as gyroptychiid osteolepids, and a phlyctaeniid arthrodire (Denisalloslells; Fig. 13K). The antiarch Sherbonaspis is similar to various Middle Devonian asterolepid antiarchs from the Nmihem Hemisphere; the genus is also recorded fi'om Kazakhstan (pante1eyev, 1993). Of biogeographic significance is the association of thelodont agnathans (Turinia cf. T. hUlkensis) with the bothriolepid antiarch Manarolepis. These two groups co-occur in other East Gondwana sequences, e.g., the Antarctic Aztec fauna (young, 1988a) and the Gneudna fauna ofW:A. (Long & Trinajstic, 2000), but the association is unknown from Nmihem Hemisphere crustal blocks, where turiniids are of Early Devonian age and BOlhrialepis is typical of late Middle Devonian and younger strata. The persistence of turiniid thelodonts until the early Frasnian is characteristic of Gondwana sequences (Turner; 1997).

The well-studied Antarctic Aztec fish fauna, of assumed Givetian-Frasnian age, with some 42 taxa (young, 1988a, 1989b, 1991a; Long, 1995a; Long & Young, 1995, Hampe & Long, 1999), includes various taxa otherwise known only from SE Australia. Examples are the acanthodian eulmacanlhus (Long, 1983; Young, 1989b; Fig. 14A-C), the lungfish Howid;Plerus (Long, 1992), the sharks AnlarctilanlJlG pl'isca and Mcmurdodus (Young, 1982; Turner & Young, 1987), and canowindrid osteolepiforms (Long, 1985; Young el al., 1992; Fig. 14D-E). The palaeoniscoid osteichthyans are also closely related (Long, 1988; Young, 1991a). This is compelling evidence

215

for continental continuity betweenEAustralia and Victoria Land at that time, and the juxtaposition of Antarctica with Australia is accepted in all Gondwana reconstructions (Fig. 16). No precise equivalent of the Aztec fauna has yet been identified in E Australia. This may be due to age differences-slightly younger than the Hatchery Creek fauna, but slightly older than fish faunas ofthe volcanic Mt Howitt province of east-central Victoria (late Givetian according to Long, 1999)-01' to the presence of a cool water 'Malvinokaffric' element in the Aztec fauna, paralleling the Early Devonian brachiopod faunas of New Zealand and Antarctica (hut less apparent in the EAustralian faunas; see Boucot el al., 1969; Talent, this chapter). Thus the diversity of chondrichthyans in the Aztec fauna (Long & Young, 1995) is not minored inknownAustralian fish assemblages. Abundance of chondrichthyans and acanthodians at the expense ofplacoderms is a feature of Devonian faunas of South America and South Africa, closely associated with Malvinokaffric invertebrate assemblages of West Gondwana (Lelievre el al., 1993; Long el al., 1997; Anderson elal., 1999). This suggests some latitudinal variation in Devonian assemblages within Australasia. Possibly connected with regional difterentiation is the prevalence of some groups (e.g. phyllolepids and crested bothriolepids) in presumed intermontane lacustrinevolcanic sequences of Victoria, compared to central Australia. Nevertheless the Middle-Late Devonian phyllolepid-antiarch placoderm associations seen in the Lachlan Fold Belt occur across central and W NSW (e.g., Young, 1999) and into central Australia (HiUs, 1959; Young, 1985). The endemic phyllolepid genusPlacolepis (Fig. 13 F), first described from the EdenComerong volcanic rift in the E (Ritchie, 1984), is also known from the Harajica fauna of the Amadeus Basin; this accords with lack of wide separation ofthe tenanes of the Lachlan FoldBelt from cratonic Australia during the late Middle Devonian.

Late Devo1lian faunas. As with marine invertebrates, the pronounced Early Devonian endemism diminishes in the Middle Devonian. By Late Devonian time fish faunas are made up of significantly more widespread or cosmopolitan taxa. Three biogeographic patterns can be distinguished (Young, 1993b, 1995a; Rich & Young, 1996). Without good age control, these were formerly interpreted to indicate general Late Devonian cosmopolitanism. Gondwanan endemism, however, generally persisted through the Frasnian (Fig. 17C). The Gogo fauna (Canning Basin; Long, 1988, 1995b) is a diverse

216

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marine assemblage, but with a substantial endemic component, e.g. 23 arthrodire genera (Fig. l3A), of which only two occur elsewhere. Late turiniid thelodonts persisted into the Frasnian in the Canning and Carnarvon basins of WA (Seddon, 1969; Turner, 1997; Long & Trinajstic, 2000); phyllo1epid p1acoderms, widespread in Givetian-Frasnian assemblages in central and E Australia, are unknown in strata of this age outside Gondwana.

In the Famennian, widespread elements useful for biostratigraphy, e.g., phoebodont sharks, are known only from marine limestones (Turner, 1982,1995), whereas typical Old Red Sandstone placoderm assemblages of the Northern Hemisphere (e.g., the Phylloiepis-BofhriolepisRemigolepis-Groenlandaspis association of E Greenland), occur with the same genera in central NSW (e.g., Hervey Group; Johanson 1997, 1998; Young, 1999). As in E Greenland, tetrapods also occur (Warren & Wakefield, 1972; Campbell & Bell, 1977), providing strong evidence of land connection with Euramerica (see below). Note that the recently claimed Carboniferous isotopic age for the E Greenland tetrapod Ichfhyosfega (Hartz ef al., 1997) is refuted by the associated placoderm faunas-not mentioned by those authors.

The youngest Devonian macrovertebrate assemblage, assigned to the latest Famennian (Young & Turner, 2000), is the Grenfell fish fauna characterised by the antiarch Grenfellaspis (Ritchie ef al., 1992), the only demonstrated occurrence outside Asia of a major endemic Chinese group (the Sinolepidae of Liu & P'an, 1958; Fig. 15G-J). Phyllolepids had become extinct by this time (Fig. 17C); other elements such as large remigolepid antiarchs suppOltAsian affinities in the latest Famennian.

Devonian veJ'tebJ'ate biogeographic patterlls. Biogeographic pattems can provide evidence of connections or barriers between regions, as

Fig. 17. Conflicting data on palaeogeographic change during the Devonian Period derived from palaeomagnetism (A-B) and vertebrate distribution patterns (C). Major counter-clockwise rotation of Gondwana from Early to Late Devonian (A-B) is implied, proximity to Asia in the Early Devonian, and a wide oceanic barrier with Euramerica in the Late Devonian. Late Devonian biogeographic patterns for Gondwana (C) imply high endemism and wide separation from Asia in the Early Devonian, proximity and connection with Euramerica near the F-F boundary, and Asian connections near the DevonianCarboniferous boundary.


distinct from evidence of palaeolatitude provided by some other data sets, e.g., palaeomagnetism (Young, 1990a). The palaeogeographic disposition of cmstal blocks possibly connected with or separated from Gondwana can be illuminated by the evidence of fossil fish distributions documented from the Australasian region. Pelagic fishes, able to traverse open oceans, are of minimal biogeographic significance, but this is not the case for groups with strong provincial patterns indicating confmement by marine barriers.

During the Early-Middle Devonian, the Australasian region constituted the 'East Gondwana Province' , one of five major vertebrate provinces (including Euramerica, Siberia, Tuva and South China) defined by Young (1981). Subsequent biogeographic analysis (Young, 1990a,1993b, 1995a, 1995b) and additional taxonomic data (e.g., Janvier, 1996) have supported this provincial pattern with minor modification r.:w and N Gondwana now appear discrete; other terranes of E Asia should be included with the South China Block). The northern blocks, Laurentia and Baltica, were characterised by osteostracan/heterostracan agnathan faunas during the Late Silurian-Early Devonian, establishing their contiguity as the palaeocontinent Euramerica. Siberia was separated from Euramerica by the Uralian seaway; it was traversed during the Early Devonian by pteraspid heterostracans, providing some relative measure of the dispersal capability of this group. Of the many smaller clUstal blocks involved (Talent et al., 1987), at least the main 'Angaran' block and Tuva share with Euramerica heterostracans and osteostracans respectively, though represented by derived groups (Blieck & Janvier, 1993). The tannuaspids of the Tuva Province represent the only extra-Euramerican occurrence of osteostracans; another major endemic group of heterostracans (amphiaspids) characterises the Angaran block. Neither group is known from Kazakhstan, which instead shows some evidence of Asian or Gondwanan affinities (Young, 1993b). The Devonian positions of both Kazakhstan and Siberia derived from palaeomagnetic data require separation oflike and juxtaposition of unlike veltebrate faunas (Fig. 16). Although they crossed the narrow Uralian seaway, Devonian heterosh'acans are completely unknown from Gondwana, so the Early Devonian 'transpressional collision' between Laurentia and Gondwana suggested by Dalziel ef 01. (1994) as part of the Acadian orogeny (Fig. 16A) can be discounted. Vertebrate distribution patterns require latitudinal separation between Gondwana and Euramerica with a width greater than the Uralian seaway at that time.

217

The Tarim, South and NOlth China terranes and Indochina shared a distinctive and highly endemic ga1easpid-yunnanolepid assemblage, indicating they were already closely associated by the Late Silurian (Tong-Dzuy et al., 1996; Young & Janvier, 1999), rather than dispersed terranes scattered along the N Gondwana margin (cf Fig. 16). The Devonian connection between North and South China Blocks (Young, 1990a, 1995b; Ritchie et 01., 1992) has been maintained on other evidence (e.g., geochemical data) consistent with the occurrence of endemic Asian fish groups (galeaspid agnathans and sinolepid antiarchs) on both blocks. A major problem with the SiluroDevonian vertebrate distributions of E Asia (Young & Janvier, 1999) is that they seem more consistent with modern geography than with widely separated Palaeozoic Asian telTanes, as are required by some Gondwana dispersal models (e.g., Metcalfe, 1996). A remote location off the Pacific margin of Gondwana would provide suft1cient isolation to develop the high endemism characterising this composite Asian terrane. However, limited exchange of some elements of the respective vertebrate faunas by Early Devonian times (e.g., the microvertebrates Ligulalepis and Poracanthodes qujingensis, yujiangolepid arthrodires, and possible affmities of Neeyambaspis; Fig. 15B-D), continuing through the Middle Devonian (species groups of Bothriolepis; Fig. 15E-F), and Late Devonian (sino1epids; Fig. 15G-J), accords with increasing proximity between Asia and E Gondwana.

Three vertebrate distribution patterns (Gondwanan, Euramerican, Asian) succeed one another in time through the Late Devonian (Fig. 17C). Though physiological adaptation could explain increased dispersal capabilities for specific taxa (e.g., Burrett et 01., 1990), shared patterns among both marine invertebrates and vertebrates require a common (extrinsic) cause stich as global change in geography or climate. The oldest of these patterns (Givetian-Frasnian) is a continuation of E Gondwana endemism exemplified by diversity of phyllolepid placoderms, a group long known from Europe (Agassiz, 1844), but with a different stratigraphic disllibution to E Gondwana (Famelmian only; Young, 1974). The succeeding Euramerican biogeographic interval is characterised by the last radiation within the distinctive cephalaspid/ heterostracan fauna of the Euramerican province: the late Frasnian psammosteid heterostracans in the Baltic region. This group disappeared completely at the Frasnian-Famennian boundary (Halstead, 1987; Young, 1981, 1987a, 1990a), after which phyllolepid placoderms make a sudden appearance in Euramerican successions.

218 AAP Memoir 23 (2000)

TAXA Yea Ghin Ghin Melbourne

Baragwanathia longifolia Lang & Cookson * *

Baragwanathia sp. 1 *

Baragwanathia sp. 2 *

Baragwanathia-like taxon *

Zosterophyllum sp. 1 *



7osterophyllum sp. 4 *

~/opella australis Tims & Chambers *

Hedeia sp. 1 * *

Yeaia flexuosa Douglas *

Bythotrephis gracilis Hall *

Table 4. Silurian (Ludlow) Baragwanathia Flora association.

Similarly, it was only after phyllolepids disappeared fi'om the vertebrate succession of E Australia that similarities became apparent between E Gondwana and the Asian blocks-in the youngest of the three biogeographic patterns (?latest Famennian-Toumaisian; Fig. 17C).

Verlebrale evidel/ce for palaeogeographic chal/ge. The major changes in vertebrate distribution patterns during the Middle-Late Devonian discussed above are assumed to reflect majorreanangements of global geography. Range enlargement into Euramerica of the East Gondwana wuttagoonaspid-phyllolepid placoderm lineage accords with increasing proximity between Gondwana and Euramerica during the Frasnian, \vith continental connection via W Gondwana at or near the FrasnianFamennian bmmdaty (Young, 1990a, 1995a). This is in contrast to the palaeomagnetic reconstruction (Fig. 16B) showing a wide ocean at that time. The distribution of pre-Famennial1 phyllolepids across the northern Gondwana margin, and their complete absence from Asia, were two predictions of the dispersal model (Young, 1981, 1989a}still intact with the subsequent discovery of phyllolepids in Turkey (Janvier, 1983) and Venezuela (Young et al., in press). The subsequent appearance of sinolepid antiarchs in the Lachlan Fold Belt of E Australia accords with a latest Devonian connection with the previously isolated Asian terranes-again not clear from the reconstructions (Fig. 16B).

In summary, vertebrate distributional data imply two chronologically separate palaeogeographic realTangements on the margins of Gondwana during the Late Devonian. The

altemative palaeogeographic model derived from palaeomagnetic data involves major counterclockwise rotation of the Gondwana supercontinent during the Devonian (Fig. 17 AB). This model implies close connection between E Gondwana and Asian terranes in the Early Devonian, and a wide ocean separating Gondwana from Euramerica in the Middle--Late Devonian (Fig. 16), just the opposite of what is indicated by the biogeographic evidence of Middle Palaeozoic vertebrates. Li el al. (1993) proposed alternative Asian migration routes between Gondwana and Euramerica, but the distinctive and readily recognised phyllolepids discussed above have never been found in South or North China, Tarim, Kazakhstan or Siberia, the blocks seeming to fonn a northem 'stepping stone' route to Euramerica on the palaeomagnetic reconstmctions (Fig. 16).

Late Devo11iall extinction event. The FrasnianFamennian boundary coincides with one of the three well-documented major global extinction events that affected the marine benthos during the Late Devonian, the Upper Kellwasser Event. For Devonian fishes, diversity peaks in the Frasnian may reflect research on a few exceptional localities (Janvier, 1996: 290), whereas extinctions among continental vertebrate faunas (e.g., agnathans of Euramerica; Halstead, 1987) must be considered in relation to marked changes in veI1ebrate distributions across this boundary. For example, the phyllolepid placoderms discussed above are one of the few groups to actually show a diversity increase across the F-F boundalY (Can, 1995), consistent with theirrange enlargement into Euramerica. Talent (1984: 76)

AAP Memoir 23 (2000) 219

rrAXA 19 Mile Frend Boola Turton Cole's Spur Boola Creek

'{3aragwanathia longifolia Lang & Cookson • • • * • '{3aragwanathia sp. • f-ycopod sp. 1 • • ~edeia sp. • • ~yniophy1e *

Zosterophyllum sp. • !Zosterophyllum australianum Lang & Cookson • iSalopella australis Tims 8. Chambers • iSalopella caespitosa Tims & Chambers • pawsonites subarcuatus Tims & Chambers • Seed'1 •

Table 5. Early Devonian (pragian) Baragwanathia Flora association.

noted the lack of evidence of increased provincialism in marine faunas required by continental connection between Gondwana and Euramerica, but this too could have been obscured by extinctions across the F-F boundary. The succeeding Asia-Gondwana biotic dispersal proposed to explain the appearance of sinolepid antiarchs in latest Famennian fish faunas of E Australia is less well supported, but also involves examples ofval'ious invertebrate groups (Ritchie et al., 1992).

NON-MARINE BIOTA The database fol' Australian Devonian

lacustrine faunas is minuscule and, accordingly, of little biogeographic relevance. Limestone nodules from a mudstone-siltstone interval in the Fahy Sandstone (Pragian) ofthe Buchan area of E Victoria (Orth et al., 1995, 120-124) have produced an abundance of the conchostracan Cyzicus talenti '5 as well as ostracods (?darwinulids), smooth gastropods, crossopterygian fish, charophytes (see below) and coprolites (WatTen & Talent, 1967; Tasch, 1987). The antiquity (Pragian) of this non-marine community is of particular interest. A xiphosuran, Kasibelinurus amicorunl has been described (Pickett, 1993) from low in the mainly fluviatile MandagelY Sandstone, about 22 km W of Parkes, frOlll what may be a lacustrine environment, though a shallow marine or estuarine fauna of lingulid brachiopods and bivalves occurs lower in the sequence (Williams, 1977).

Macrofloras (J.G. Douglas) Constmed broadly, there seem to have been

three macro-floras in the eastern Gondwana

region during Devonian times: the Baragwanathia flora (late Ludlow-Pragian), less well-known Emsian-Givetian flora/floras, and a broadly Late Devonian Leptophloeum floral'. According to present data, there seems to have been little or no biogeographic differentiation in the macro-floras in the eastern Gondwana region during Devonian times, but the database is patchy, and is almost non-existent for the EmsianGivetian interval.

The Baragwanathia flora seems to have persisted without major change for at least 20 million years and to have been widespread. The key taxon, Baragwanathia longiJolia, extended beyond eastern Gondwana to Laurentia, from whence Hueber (1983) described B. abitibiensis from the Abitibi River, Ontario, Canada, in the Sextant Fonnation of middle-late Emsian age. This correlates approximately with the youngest occunences of the Baragwanathia flora in SE Australia. The record of the latter is based primarily on specimens, often fragmentmy, from offshore flyschoid environments 17,

Late Devonian floras have been repOlied from numerous molasse basins in E Australia from E Victoria to NE Queensland. A remarkable predominance of Leptophloeum may reflect prominence of fine-grained sandstones, the host for the comparatively large casts comprising the bulk of the available fluvial specimens. As far as can be discerned, these assemblages with equivalents in other areas of eastelTI Gondwana do not display any biogeographic differentiation.

Much less well known assemblages suggest the possibility of a third macroflora. An early Eifelian assemblage from the Storm Hill Sandstone of the Broken River area of NE

220 AAP Memoir 23 (2000)

Locality Psilopsida Lycopsida Pteridophyta Sphenopsida Cordaitales

Bindaree Unident. sp. Lycophyte sp.

Mansfield Lepidodendron mansfieldense

Avon River Lepidodendron austra/e

Freestone Taeniocrada Lepidodendron Unidentified sp.1 Cordaites Creek langi sp. australis Iguana Creek Archaeopteris

howitt( Sphenopteris iguanensis

Tabberabbera Archaeopteris Aterocalamites sp., sp., Phyllotheca Rhacopteris sp., Unidentified sp., sp.2 Sphenopteris sp.

Combienbar Lepidodendron sp.

Genoa River Barinophyton Archaeopteris Unident. sp.3, Cordaites citrullifome howitt( Unident. spA australis

Spenopter/s camei

Table 6. Plant assemblages, Late Devonian-Carboniferous, SE Australia.

Queensland, although with some Leptophloeum assemblage affiliation, may also represent a third macroflora. Some Leptophloeum assemblages from NE Australia may be Early Carboniferous, rather than Late Devonian.

Early-Middle Devolliall. The record of plant life fi'om the Early and Middle Devonian of Australia is based primarily on specimens from offshore flyschoid environments in the SE ofthe continent. Most interest has focused on the Baragwanathia flora, dominated by B. longifolia Lang & Cookson. These floras, long taken as representative of the first major terrestrial cover, consist of a diverse assemblage including relatively large species with well-developed vascular tissue. Palynological evidence however shows that the progenitors, the first land plants, are known from membrane-bound spore tetrads of Ordovician (early Llanvirn) age (Gray & Boucot, 1977; Kemick & Crane, 1997), rather than fl.·om macro-remains.

Couper (1965) demonstrated that in the key Yea area ofthe Melboume Trough there were two clearly defined plant-graptolite horizons separated by about 2000 m of strata IS. A substantial

difference in age between the two horizons was indicated when a diverse Baragwanathia Flora assemblage from Couper's "Lowel; or No.1 PlantGraptolite Horizon" at Limestone Road, was found associated with Bohemograptus bohemicus, indicative of Late Silurian (Ludlow) age (Garratt & Rickards, 1984) (Table 4). The 'Upper, or No.2 Plant-Graptolite Horizon', about 2000 m stratigraphically higher, bears essentially the same flora as the type locality of BaraglVanathia longifolia at the 19-mile quany on the Warburton-Woods Point road (Table 5).

Major contributions on the floras of both plantgraptolite horizons were presented byTims (1980) and Tims & Chambers (1984). Tims confirmed allocation of Baragwanathia to the Lycophytina, and considerably enlarged knowledge of the Silurian flora from Yea, as well as the younger (Pragian) occurrences on the basis of new collections from Frenchmans Spur, Cole's Clearing and Boola QualT)' (Table 5). Three new taxa were described by Tims & Chambers (1984). Tims (1980) considered that B. longifolia from the 'Lower Plant-Graptolite Horizon' displays differences in morphology from topotype B. longifolia-a development not unexpected


considering the assemblages represent plants that lived ten million years or lllore apart.

In SE Australia B. longifolia has also been obtained from the Mullamuddy Formation (Pragian?) from near Mudgee; other taxa of the Baragwanathia Flora have been noted in the Mt. Daubeny FOlmation, NE of Broken Hill, NSW; it contains 'Psilophyton sp.' (Neef et al., 1989), regarded as indicating an Early Devonian (Lochkovian) age.

The undescribed Turtons Creek assemblage (Table 5), thought to be Pragian, contains specimens considerably less weathered than any others so far collected; it has potential to provide material for cuticular and other anatomical study.

Late Devonian floras have been repOlted fi'om numerous molasse basins in E Australia from E Victmia to NE Queensland. These include localities known for well over a century, dominated by Leptophloellm australe McCoy. Species described over forty years before any members of the much better known Early DevonianBaragll'anathia Flora include Barinophytoll obscurum, Archaeopteris hOlVitti, and Calamitales from the Genoa Riverjust nOlth of the NSW-Victoria border (Dun, 1897) (Table 6). Other salient occurrences recorded by McCoy (1876) and Douglas (1959, 1960) areli-om Bushy Park (Avon River), Tabberabbera, and Freestone Creek, Victoria, the last two with, respectively, the psilophyte Taeniocrada langi (Stockmans), and Archaeopteris howitti McCoy (Table 6). An unnamed Iycophyte (White, 1986) from Bindaree, VictOlia, appeat~ to be in the lineage from large woody Baragwanathia to herbaceous lycophytes. Small herbaceous plants were already well advanced. I'

Among numerous localities in NSW are ocCIUTences in the Eden-Merimbula, Wellington and Mudgee areas, and in the Bundock Creek Group of the Broken River region and the Drummond Group ofNE and central Queensland respectively. None of these has been the subject of intensive or recent investigation.

Palynofloras (G. Playford) Terrestrial and marine palynomorphs - spores

and organic-walled microphytoplankton respectively - have been described or recorded fl'om Devonian (especially Givetian-Frasnian and latest Famennian) strata of several Australian sedimentary basins. The plant microfossils have been retrieved entirely from subsurface samples insofar as surface exposures are nOl1nally too highly weathered to sustain palynomorph preservation (e.g., Playford, 1985, p. 248).

Spores. Australian spore floras of possibly Early Devonian age are known thus far only from the

221

Adavale Basin, Queensland (Price, 1980; Hashemi, 1997), in patticular from the basal unit therein (Eastwood Beds: ?Emsian). This contains, inter alia, abundant tricrassate/apiculate forms (e.g., Rotaspora, Ti"icidarisporites) and is devoid of ancyrate spores. Only broad morphological similarities are evident with coeval Northern Hemisphere palynofloras.

Stratigraphically higher,Australian Middle and Upper Devonian (Givetian-Frasnian and possibly early Famennian) assemblages likewise include many endemic taxa, but an increasing cosmopolitanism is attested by their content of such fmIDs as Geminospora /emurata Baime emend. Playford, 1983, Verrllcosisporites scurrus (Naumova) McGregor & Camfield, Teichertospora torquata (Higgs) McGregor & Playford, Cristatisporites Iriangulatus (Allen) McGregor & Camfield, Ancyrospora langii (Eisenack) Richardson, Emphanisporites annu/atus McGregor, Cymbosporites magnijicus (McGregor) McGregor & Camfield, and Rhabdosporites langii (Eisenack) Richardson. Indeed, McGregor & Playford (1993) were able, on the basis of shared specific taxa, morphons, and gross morphological features, to cOlTelate Australian strata-pdncipally of the Canning, Carnarvon, Amadeus, and Tasmania Basins~variously with the spore zones of the Euramericau Old Red Sandstone Continent (ORSC; Richardson & McGregor, 1986): see McGregor & Playford (1993, p. 39) for infelTed Australian occun·ences of, in particular, the optivustriangillatlls, ovalis-bulliferus, and torquatagracilis spore zones. McGregor & Playford (1993) thus demonstrated similarities between Euramerican and Australian spore assemblages suft1cient to indicate palaeo-geographic proximity, and concomitant floristic interchange, between Eastem Gondwana and the ORSC during mid to Late Devonian time. More recently, Streel & Loboziak (1996, p. 583) cOlmnented that, fl'om McGregor & Playford's data, some 150 spore species appear to be endemic to Australian Middle to lower Upper Devonian sequences, suggesting that "Australia might well cOl1"espond to a single phytogeographic entity" within or discrete from the overall Western Gondwana-Southern Euramerica Province; see also Balme (1988, pp. 109, 152).

An absence or dearth of palynological data currently exists for the lower to middle Famennian, in Australia and in many other regions. However, in general, uppermost Famennian assemblages are well documented. For instance, in the Canning and Bonaparte Basins ofNW Australia, abundant and diverse suites are characterized, as globally elsewhere, by the distinctive cavate/reticulate species Retispora

222

lepidophyta (Kedo) Playford, widely accepted as being restricted to the latest Famennian ('Strunian') (e.g., Playford, 1993; Playford & McGregor, 1993). Other notable cosmopolitan or near-cosmopolitan components afthe Austra1ian lepidophyta-nitidus zonal representation (see McGregor & Playford, 1993) include Verrucosisporites nitidus Playford, Auroraspora macra Sullivan,Indolriradites e.,plaJlatus (Luber) Playford, and TlI1I1ulispora rarituberculata (Luber) Playford. However, as with older Australian Devonian palynofloras, the lepidophyta assemblage includes a significant proportion of seemingly endemic species, e.g., Granlilatisporites jrllstulentlls Bahne & Hassell emend. Playford, Apiculatisporis mOl'bOSliS

Bahne & Hassell, Reticuiatisporites ancoralis Balme & Hassell, and Brochotriletes textilis (Balme & Hassell) Playford (see Balme & Hassell, 1962; Playford, 1976).

The trend towards accentuated spore cosmopolitanism enhances the perception that an appreciable degree of commonality in the composition of the world's land vegetation was attained in later Devonian time (e.g., Edwards, 1973). In many instances, botanical affinities of Devonian dispersed spores remain highly speCUlative (Bahne, 1995), but they certainly represent the products oflargely coastal lowland communities that included progymnosperms (notably archaeopterids), zosterophyllophytes, rhyniophytes, lycopods, and pterophytes.

Orgallic-walled microphylopiallkloll. Lower Upper Devonian marine strata of the Camarvon and Canning Basins, WestelTI Australia, have yielded profuse assemblages of microphytoplankton (acritarchs and prasinophyte phycomata) in association with the telTestrially sourced spore floras discussed above. The Carnarvon Basin phytoplankton, from the early Frasnian Gneudna Formation (Playford & Dring, 1981; Playford, 1981), exhibit a distinctive provincial character, as noted by the latter authors and subsequently by Wicander & Playford (1985) and Colbath (1990a). The coeval phytoplankton suite from the Limestone Billy Hills Reef Complex, N Canning Basin (Colbath, 1990b), signifies a greater diversity and perhaps less provincialism, but it includes, not unexpectedly, many Gneudna forms newly instituted or recorded by Playford (in Playford & Dring, 1981); e.g., Dictyotidillm pro/aftoJl, Lomat%pas cellulosa, Papulogabata allnulata and Melikeriopalla veJ/liiosum. Both Western Australian Frasnian suites do contain appreciable proportions of cosmopolitan or nearcosmopolitan species; e.g., Maranhites brasiliensis Brito, Polyedryxium pharaonis


Deunff, Cymatiosphaera perimembrana Staplin, Daillydium pentaster (Staplin) Playford, Multiplicisphaeridium ramispinosum Staplin, Unellium lunatum (Stockmans & Williere) Eisenack, Cramer & Diez, U piriforme Rauscher, and Solisphaeridiu1I1 spillog/obosllm (Staplin) Wicander. Many oftllese are also shared with the palynoflora ofthe lower Shishtu Fonnation oftlle Central Iran Basin (Hashemi & Playford, 1998) in accord with the inferred northern Gondwanan palaeoproximity of these regions in Late Devonian time. Colbath (I 990a) considered that his Canning Basin palynoflora evinced greater similarity with the Frasnian assemblage of Iowa (Wieander & Playford, 1985) than with those of other Gondwana regions. In general, it should be emphasized that attempting to establish the biogeographic affinities of Australian Frasnian phytoplankton suites is hindered by deficiencies in the number of adequately documented assemblages from well-dated strata elsewhere (with the exception ofW Canada, Midcontinent U.S.A., and Belgium: Wieander, 1996, p. 512).

Attenuated prasinophyte and acritarch productivity and diversity are known globally to characterize the latest Devonian and Carboniferous, and this is exemplified by the depauperate phytoplankton component of the palynoflora described by Playford (1976) from the Fairfield Group (latest Famennian-Toumaisian) of the N Canning Basin. Among the relatively few forms recorded, the majority are known to be widely distributed geographically in strata contiguous with the Devonian-Carboniferous boundary (Playford, 1993; Playford & McGregor, 1993); e.g., Gorgonisphaeridium absitum Wicander, G. ohioense (Winslow) Wicander, Stelliniu1l1 micropolygonale (Stockmans & Williere) Playford, and Tornacia sarjeantii Stockmans & Williere. These impatt a distinctly cosmopolitan complexion to the Fairfield assemblage.

Charophyta (M. Feist) Charophyta are now known from two

Devonian horizons: the lacuslline FallY Sandstone (late Pragian or a little earlier) ofE Victoria (Feist et al., unpub. data), and the marine Mytton Formation (seemingly early Frasnian) of the Broken River region ofNE Queensland (Feist et ai., 2000). Recent discoveries of charophytes from the Faity Sandstone (no younger than Pragian) high in the Snowy River Volcanics ofE Victoria has demonstrated that Charophyta occulTed in Eastem Gondwana early in their histOlY.

The Pragian charophytes from the Fairy Sandstone have been obtained from limestone nodules in an interval of tuffaceous siltstones and shales where they occur in association with fish


remains, branchiopods (Cyzicus talel/Ii Tasch), nondescript ostracods and gastropods presumed to be non-marine forms, and plant remains (WalTen & Talent, 1967; Tasch, 1987).

The Frasnian charophytes (Feist et al., 2000) were obtained from the Mytton FOlmation in the Page Creek area of the Broken River region of NE Queensland, a few metres above the ently of Icriodus symmetricus at the top of the Stanley Limestone Member of the Papilio Formation (Mawson & Talent, 1989)'". The charophytes occur in mudstones/muddy siltstones associated with a marine fauna consisting of an undescribed fauna of brachiopods (especially atrypids, dalmanellids and pentamerids), tabulate corals, (especially Alveolites) and rugosans. The best approximation to age is early Frasnian.

In the absence of data from other localities, these occurrences do not have special relevance for palaeobiogeographic questions, though their generalised nature, and the conspecificity of the Frasnian form with the widely-distributed Sycidium reticulatum Sandberger is consistent with ease of distribution of charophytes between Baltica and East Gondwana during the Devonian.

ACKNOWLEDGEMENTS The coordinators (JAT & RM) are grateful to

the many authors who produced manuscripts on time and, without losing a sense of humour, reconciled themselves to what proved to be an inordinately protracted process of editing, to Alison Basden for making available, at a critical time, her manuscript bibliography of Australian Devonian fish, to Kath Grey for providing a copy of her unpublished (1984) compilation of ranges of Canning Basin Devonian brachiopods in relation to conodont zones then in use (the basis of Table 3), to Scott Brownlaw for allowing use of material on Western Australian rugose coral faunas fl'om his PhD studies, to Sue Turner for critically reading a large part of the text, and to Tony Wright for kindly undertaking a careful reading of our entire manuscript in pursuit of infelicities. The drafting was by our friend Dean Oliver.

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Aitchison, J.C., 1988. Radiolaria from the southern part of the New England Orogen. 49-60 in Kleeman, J.D. (ed.), New England Orogen tectonics and metallogenesis. University of New England, Annidale.

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Aitchison, J.C., 1993a. Albaillellaria from the New England orogen, eastern NS W, Australia. Marine Micropaleontology 15, 353-368.

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Banks, M.R. & Rickards, R.B., 1989. Early Devonian graptolites from Tasmania. Papers alld Proceedings a/the Royal Society of Tasmania 123, 111-117.

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APPENDIX 1: ENDNOTES Introduction and Conodonta (R. Mawson & 1.A. Talent)

lThe primary basis for stratigraphic alignments in E Australia has been conodont data (Fig. 5; Appendix 2). Conodont faunas from E Australia span a large part of the Early Devonian. Small faunas with Icriodus woschmidti from the Ehnside Formation of the YassBowning area of NSW (Link & Druce, 1972), from the Clandulla Limestone of the Rylestone-Cudgegong area of east-central NSW (Colquhoun, 1995),fi'om the uppennost Jack Fonnation ofthe Broken River region of NE Queensland (Simpson, 1995a, 1995b), and a small fauna including Ozarkodina excavata excavata and a morph of O. remscheidensis remscheidensis from the \Vombat Creek Group at Pyle's limestone deposit near Benambra (Whitelaw, 1954, Fig. 3D; Simpson & Talent, 1995; Talent et al., unpub. data) indicate that earliest Lochkovian horizons occur in all ofthese areas but virtually no taxonomic data are available for associated shelly faunas from these earliest Devonian horizons. Better known shelly faunas, but less well dated, are known from several latest Silurian--earliest Devonian sequences in SE Australia.

Conodonts from the Lilydale Limestone (Wan et al., 1995) show that sequence to span at least the kindlei and pireneae zones of the Pragian. Conodonts from a carbonate fan in the Tyers-Boola area in the \Valhalla Synclinorium, 80 km E ofMelboume, span an interval from within the sulcatus Zone through the kindlei Zone, and presumably pireneae Zone, into the early Emsian dehiscens Zone (Mawson et aI., 1992; Mawson & Talent, 1994a; Mawson, 1997). To the north in the Walhalla Synclinorium, limestone clasts from debris flows and isolated limestone olistoliths suggest cannibalisation of a persistent carbonate platfonn that once fringed a now buried 'lost terrain' in south~vest Gippsland. The precise ages of these have not been detennined relative to the Lochkovian-Pragian boundary (Mawson & Talent, 1994a). Conodont data from limestone olistoliths at Loyola, NE of Melbourne (Mawson et al., 1992) are also dated as Pragian; the source of these is uncertain. South of the 'buried terrain', along the W flank of Waratah Bay, three Devonian units crop out: the Waratah and Bell Point Limestones-Pragian, sulcatus, kindlei and possibly pireneae zones (Talent, 1989; Bischoff &


Argent, 1990; Mawson et al., 1992; Mawson & Talent, 1994a)-and the flyschoid Liptrap Fonnation, possibly an extension of the Walhalla Group of the Walhalla Synclinorium to the N.

Several carbonate horizons occur in the sometimes richly fossiliferous Wentworth Group, extending through Tabberabbera (Talent, 1963), NW of Bairnsdale. Conodonts from these outcrops and from limestone clasts from the oldest unit in the \Ventworth Group, the Wild Horse Fonnation, show it to be no older than late Pragian Polygnathus pireneae Zone (Mawson et al., 1992). The brachiopod faunas, principally from the Kilgower Member of the Tabberabbera Formation are therefore construed as being approximately dehiscens Zone and thus approximately coeval with faunas oftheBuchan Caves Limestone, Cavan Formation and perhaps younger units of the Buchan and Murrumbidgee Groups.

Conodonts from the Buchan Group of E Victoria have demonstrated that the group spans most of Ems ian time (dehiscells to sera/intis zones) and apparently much ofthepirelleae Zone as well (Mawson, 1987a; Mawson e/ al., 1992). The Taravale Fonnation has produced important dacryoconarid faunas and late dehiscens Zone and perboflus Zone ammonoids and dacryoconarids (Erben, 1964, 1965; Mawson, 1987a; Alberti, 1993, 1995, 1999). The presence of strata aligning with the pirelleae Zone low in the Buchan Caves Limestone at Buchan and Bindi is inferred from the widespreadpimneae Zone transgression identified in analogous sedimentary sequences elsewhere in SE Australia, most notably in the Cavan Fonnation ofSE NSW (Mawson e/ a/., 1992), but this interval may predate the well-preserved brachiopod faunas of the Buchan Group (Talent, 1956a; Valeutine, 2000 liS.).

Graptolithina (R.B. Rickards &A.J. Wright) 2Graptoloids occur in Australia in Lochkovian and

Pragian sequences. Monograptus ulliformis lInifarmis has been recorded from the Lochkovian at Limekilns, NSW. The Pragian species, M. thomasi has been recorded from the Wilsons Creek Shale, Victoria, and Flowery Gully, Tasmania,M. aequabilisnotoaequabilis from the 20-mile QUarry, Victoria, and M aequabilis cf.notaaequabilis from the Mathinna Beds, Tasmania. Because graptoloids were planktonic, it is not surprising that these show little provinciality.

In their paper concerning graptoloids from Limekilns, NSW, Rickards & Wright (2000) reviewed the Australian Devonian graptoloids. The text which follows is taken from their paper. Jaeger (1978, p. 505) noted that material from Cheeseman's Creek near Orange, NSW was "reminiscent of M. uni/ormis". He subsequently (Jaeger, 1988, p. 431) stated that "there are no indubitable fmds of Lochkovian graptolites in Australia", and (Jaeger, 1988, Fig. 1) showed the Cheeseman's Creek occurrence as questionable M. lIniformis; he did not comment on this in his text.

247

Sherwin (1979) listed this Cheeseman's Creekfonn as M. aff. thomasi Jaeger, 1966, a Pragian species. However, were this correct, then the graptolites occurring with "M. aff. thomasi" at Cheeseman's Creek are unlikely to be the Pridoli species, M. transgrediens Pemer, 1899 (Jaeger, 1978, p. 505; Sherwin, 1979, p. 161). Two specimens from Cheeseman's Creek are Monograptus prognatus Koren (Koren, 1983), aPridoli species (Rickards & Wright, 2000, Fig. 2A-C); and M transgrediens Perner, 1899. This locality must therefore be Pridoli (latest Silurian) and not Devonian.

Rickards & Garratt (1990) recorded M. aff. uni/armis angustidens Pribyl, 1940, from high in the Humevale Siltstone of Victoria, but considered it more likely to be late Pridoli than Lochkovian. Jenkins (1982) also convincingly argued a late Pridoli age for his record of M. cf. angustidens from the Elmside Fonnation of Yass, NSW; although this fonn has been revised by Rickards & Wright (1999) who described it as Monograptus hornyi Jaeger, 1986, suggesting a late Pridoli age. Thus the only Lochkov graptoloids recorded with certainty from Australia are: M, u. Ul1ifol7l1is Pribyl, 1940 described by Rickards & Wright (2000) from the Limekilns Fonnation at Limekilns, previously identified asM. yukonensis by Packham (in strusz e/ ai., 1972); andM aequabi/is (pribyl, 1941) described by Jaeger (1966) from Victoria.

Jaeger (1966, 1978, 1988) also reviewed the Australian Pragian graptolites listingM thomasi Jaeger, M aequabilis llotoaequabilis Jaeger & Stein, and M. yukanensis Jackson & Lenz. Jaeger (1978,p. 506) fIrst of all listed Packham's (in Strusz e/ ai., 1972) identification as M. cf. yukonensis, but he later (corrected by Jaeger's hand inreprints) deleted the 'cf.'. However, in his lists (1978, p. 502, Table 1) he recorded M)~ yukonensis only with an interrogative. This fonn has been reidentified as M. u. uniformis (see above).

Other Pragian graptolites recorded by Rickards & Banks (1979) and Banks & Rickards (1989) include: M. aequabilis cf notoaequabilis Jaeger & Stein from the Mathinna Beds of Tasmania, by the fonner; and M. thomasi Jaeger from Flowery Gully in Tasmania by the latter. The fIrst ofthese records was not mentioned by Jaeger (1988). The only other Devonian graptolite in Australia appears to be Dendrograptus sp. recorded in Rickards & Wright (2000, Fig. 2D).

Brachiopoda (J.A. Talent) 3 Australocoelia is also known from Early Devonian

(?Lochkovian) clasts in Permian glacials from the Ovens valley ofNE Victoria (Talent, unpub. data). Their derivation is problematic, though associated clasts have produced the rhynchonellidine Notoconchidiwn Gill; the latter could have been derived from the HeathcoteRedcastIe area of central Victoria where N. tasmaniense (Etheridge) is found in Unit 2, the Notoconchidiu111 Beds or Dealba Member (Edwards e/ ai., 1998),ofthe Mt Ida Fonnation. N. tasmaniense (Etheridge), first

248

recorded fiolll the Wynyard Tillite of Tasmania, is a senior synonym of N. jlorenceJ1sis Gill from \V Tasmania and N. thomasi Gill from the Mt Ida Fonnation. Elements of the same fauna are found in allochthonous blocks from Pennian glacials in South Australia and from Pemlian glacials reworked into Cretaceous sediments at White Cliffs in \VNSW (Dun, 1898; Giirich, 1901; Campbell et al., 1977; Flint et al., 1980).

"The Silurian and especially Early Devonian sequences of central and E Victoria tend to be dominated by brachiopods and, in carbonates, by coralbrachiopod faunas. Some of the brachiopod faunas have been monographed, but most have still to be described and evaluated for their chronologie and palaeoecologic significance. Some of the brachiopod faunas have chronologie control from associated graptolite or conodont data, but there is a lack of tight chronologie control for the late Ludlow to end-Lochkovian. With the salient exceptions of the diverse brachiopod faunas from the Buchan Group (Talent, 1956a; Campbell & Talent, 1967; Valentine, 2000) and small faunas from the Coopers Creek Limestone (Philip, 1962), the \Varatah, Bell Point and Lilydale Limestones (Talent, unpub. data), and from limestone olistoliths in the Walhalla Synclinorium (Talent, 1956b), preservation of most faunas as moulds has acted as a disincentive to taxonomic study. Seven of the ten Ludlow to Emsian brachiopod-based faunal units proposed by GalTatt & Wright (1988) were based on Victorian faunas. Important in this synthesis were notanopliids (Garratt, 1980, 1983a, 1983b); these tend to be more frequently encountered in arguably offshore, slope facies into which many of them may have been transported attached to nekton (Gratsianova & Shishkina, 1977).

Chronologic alignments in the thick clastic late Ludlow-Lochkovian sequence of the HeathcoteRedcastle-Costerfield area are problematic, but data from poorly preserved trilobite faunas (Holloway & Neill, 1982; A. Sandford, pers. comrn.) provide some improvement in ages relative to the Silurian-Devonian boundalY. Latest views on the Mcivor Sandstonerecently expanded to include the former units 3 and 4 of the Dargile Formation in the type HeathcoteRedcastle area (Edwards et al., 1998}-are that it may be entirely late Silurian.

Though chronologie alignments are imprecise, a broad biofacies pattern {Talent, 1964; unpub. data} is apparent for the Mclvor-Mt Ida interval, especially for the latter ifviewed in a generalised W-E sequence from Heathcote-Redcastle towards the \Valhalla Synclinorium, through Eildon and Jamieson (Talent, unpub. data from collections by -Whitelaw and Bell). Nearshore faunas dominated by bivalves tend to be replaced laterally and up-sequence by faunas dominated by rhynchonellidine and retziidine brachiopods with occasional 'Howellella " replaced in turn (presumably downslope) by faunas dominated by dalmane1lidines


such as Isol'this, plectambonitaceans, leptaenids, orthotetaceans, pentameridines, atrypidines including Austratina and, rarely, larger spiriferidines. Brachiopods in this last biofacies tend to occur rarely in generally thin intervals of comminuted shell 'chaff' with associated crinoid fragments and rare solitary rugose corals, and, infrequently, trilobites and molluscs-rarely complete and thus seldom identifiable-testifYing to comminution during and perhaps prior to major downslope transpOlt. Tending to be farther offshore are predominantly pelagic faunas with graptolites, dacryoconarids, occasional bivalves (Panenka, Hel'cynella), cephalopods and, rarely, conulariids, algae and land plants. These rarely fossiliferous slope environments do, however, have rare unbroken shells: notanopliids, small chonetidines and lingulaceans, perhaps autochthonous, but conceivably transported offshore attached to nekton.

'The Late Silurian Clonbinane (~ Mt Philippa) Sandstone Member and the 'Macropleura Band', respectively at, and about 130 m above, the base of the Humevale Siltstone, have produced diverse faunas (Williams, 1964; Garratt, 1983b), but these are indubitably Late Silurian in age and, moreover, have not been monographed.

6Two Pragian limestone units, the Waratah and Bell Point Limestones, separated by an angular unconfonnity, outcrop on the \V side of\Varatah Bay; ages of these are constrained by conodont data to being respectively early Pragian sulcatus Zone and late Pragian pil'eneae Zone extending perhaps into earliest Emsian dehiscens Zone (Talent, 1989; Bischoff & Argent, 1990; Mawson et al., 1992; Mawson & Talent, 1994a).

7A faunule of Middle Devonian brachiopods of possibly early Givetian age (de Jersey, 1966) has been described by McKellar (1966a, I 966b) from bore cores from the Etonvale Fonnation in the subsurface Adavale Basin ofS\V Queensland. McKellar believed them to be Eifelian in age, but de Jersey (1966) inferred from palynology that they were early Givetian. The faunule contains the following forms: a new genus and species of atrypidine, Neocoelia boucoti (McKellar), a nondescript, crushed l'eticulariid, Undispirifer? spinatus McKellar, and a crushed centronellidine described as R11ipidothyris go-wanensis McKellar. The age and biogeographic implications ofthis faunule are problematic.

sThe brachiopod-bearing horizons of the Canning Basin tend not to yield good conodont data and, moreover, collecting of the brachiopods pre-dated most of the conodont work. Ranges of the brachiopods relative to the conodont zonation are therefore imprecise, occasionally conjectural. The stratigraphic units recognised in the Canning Basin are faciesrelated; most extend through several conodont zones: the Pillara Limestone extending from Givetianpossibly as old as Middle varclis Subzone-through


the Frasnian until late Famennian, the Napier Fonnation and Windjana Limestone from perhaps late Givetian through to late Famennian. Combining the faunas ii-om such long-ranging units is of doubtful value because they may span several brachiopod 'zones'. Attention is drawn to the meticulous work by Strusz (1992) who provided precise locations and stratigraphic positions for all Devonian brachiopods from the Canning Basin.

9Because of a dearth of carbonate horizons, only a modicum of chronologie precision is available from Late Devonian conodonts of E Australia (Mawson & Talent, 2000). The exception is the Burdekin Basin where major marine transgressions during the Famennian occurred in the margil1ifera and expansa zones (Mawson & Talent, 1997). One widespread transgression extended over much of E NSW during the Frasnian; this has been dated as latest Frasnian linguiformis Zone at Ettrema, NS\V (pickett, 1972). An early Famennian age has been suggested recently (Jones et al., 2000) for a subsequent, less extensive transgression into west-central NS\V.

Radiolaria (I. Aitchison) lOWell-preserved radiolarians were first identified

from the Tamworth Belt (= Gamilaroi Terrane) ofE Australia, particularly in the vicinity of Tamworth, at the end oflast century (David & Howchin, 1896; David & Pittman, 1899; Hinde, 1899; Aitchison & Stratford, 1997; Mawson et al., 1997). Recent work has confumed the presence of abundant Devonian radiolarian faunas at numerous other localities in the same region, especially in tuffaceous sediments (Aitchison, 1988, 1993a; Aitchison & Flood, 1992, 1994; Aitchison ef al., 1992; Ishiga, 1989; Ishiga & Leitch, 1988; Ishiga ef al., 1987, 1988; Spiller, 1992; Dongal, 1995; Stratford, 1995; Stratford &Aitchison, 1997; Metcalfe et al., 1997). These faunas are often not well preserved and generally comprise only radiolarians; this makes precise alignment with biozonal schemes based on other groups, notably conodonts (cf. Mawson ef aI., 1997), difficult. Stratford & Aitchison (1997) suggested a late Early Devonian to early Middle Devonian biostratigraphy for the Tamworth terrane with seven radiolarian assemblages (Aitchisonet aI" 1999). These, in ascending order, are: the Stigmosphaerostyla honida, Helenifore laticlavium, Circulaforma admissarills, Helenifore pilosidiscus, Protoholoeciscus hindea, Ceratoikiscum regalinodus and Trilonche mina."\' assemblages.

Foraminiferida (K.N. Bell) IlWhen comparing Emsian foraminiferal faunas of

Australia, relevant data on faunas are available for the USA (Cushman & Stainbrook, 1944, Ireland, 1939; Stewart & Lampe, 1947; Sununerson, 1958), Poland (Malec, 1992) and Germany (Bartenstein, 1937; Langer, 1969). A large number of studies have focused on foraminiferal faunas from the EifeHan in the

249

Northern Hemisphere (e.g., Blumenstengal, 1961; Bykova, 1955; Duzynska, 1956, 1959; Sobat, 1966; Toomey, 1968; Eickhoft~ 1968, 1970; Pichier, 1971; Malec, 1984, 1992; Malec & Studencki, 1988). This seems to be an adequate database against which to consider the level of endemism presented by Australian Emsian-Eifelian faunas.

Stromatoporoidea (B.D. Webby) 12The fIrst records of Australasian stromatoporoids

are those of Etheridge (1895, 1917a, 1918, 1921a), and Chapman (1912a, 1912b, 1914a, 1914c) from the Siluro-Devonian successions of NSW and \Vestern Australia, and of Victoria, respectively. The next most impOliant contribution are the works by Ripper (1933, 1937a, 1937b, 1937c, 1937d, 1938). Since then, several occurrences have been documented from the Broken River region ofNE Queensland (Mallett, 1968,1970., 1970b, 1971), the Yass area of central NSW (Birkhead, 1978), the Recfton Group of New Zealand (Cockbain, 1965) and from the Canning and Carnarvon basins of WestemAustralia(Cockbain, 1975, 1984, 1985, 1989). Recent work by Webby ef al. (1993), Webby & Zhen (1993,1997, in prep) and Cook (1999) have provided further documentation of the Victorian, central NS\V (Limekilns) and N Queensland (Broken River and Burdekin Basin) faunas.

Ostracod. (p.I. Jones) i3Most Devonian ostracods appear to have been

benthic, although some widely distributed groups such as the entomozoids, are generally thought to have been pelagic. The distribution of the benthic forms, like those of brachiopods and corals, was controlled by environmental factors such as salinity, temperature, substrate and water depth. Although detailed sampling is necessary before generalisations can be made about the ostracod content of a given area (Reynolds, 1978), zoogeographic comparisons and contrasts can be made of similar biotopes in different areas, both on local and intercontinental scales. Of the several schemes of Devonian ostracod biotopes taken to be related to increasing water depth from high-energy to low-energy environments (polenova, 1971; Becker, 1975, Becker & Bless, 1990, Berdan, 1990; Wang, 1988), Wang's (1988), fIve Palaeozoic ostracod associations (leperditiid, palaeocopid, smooth-podocopid, spinose-podocopid, and entomozoacean associations) are used in this account. Only the leperditiid association, indicative of very shallow-water tidal flat, lagoonal or deltaic environments and brackish or hypersaline waters (Berdan, 1984, 1990; Wang, 1988) is missing in the Devonian. The sole occurrence of this association in Australia is in the Late Silurian (early Ludlow) Cliftonwood Limestone at Yass, NSW (Chapman, 1909). TIle entomozoacean association is represented by mainly cosmopolitan species in the Late Devonian of the Bonaparte and Canning basins of West em Austra lia.

250

Vertebrata (G.C. Young, J. Long & C. Burrow) 14Earlyllower vertebrates first appear in the fossil

record in the Late Cambrian-Early Ordovician, and by the Middle Palaeozoic had diversified into some 25 orders (for a comprehensive, authoritative account see Janvier, 1996). The two major groupings offish-like vertebrates are the jawless agnathans, and those with the anterior gill arch skeleton modified into a biting apparatus (the jawed Of gnathostome fishes). Together, these groups apparently occupied almost all the habitable aquatic environments of the Early-:Middle Palaeozoic. The DevonianPeriod, the 'Age of Fishes' , preserves the first major radiation of the jawed vertebrates, and provides a better fish fossil record than other parts of the geological column. This may have been due to lack of competition from the terrestrial realm, or to better preservation and chance of discovery of heavily ossified skeletons, the primitive condition retained by most Silurian-Devonian fishes. The Devonian radiation of gnathostomes into nOlHnarine aquatic environments may have been in response to increased productivity resulting from establishment of a terrestrial flora and fauna by the end of the Silurian (e.g., Gray, 1988; Jeram ef al., 1990).

The Australasian region has yielded abundant and diverse Devonian fish faunas, although only in the last 25 years has detailed study provided data relevant for biogeographic analysis. Before this, a bias towards cosmopolitan or widespread taxa, because of the tendency to initially identify familiar fonns innew areas (Young, 1995a), led to a general impression (e.g., Hills, 1958) that the Devonian fish faunas of Australia were essentially similar to those from the 'Old Red Sandstone' deposits of the Northern Hemisphere.

Devonian vertebrate fossils from Australasia were fITStreported from Buchan (Victoria) by McCoy (1876). Subsequent descriptions of the Taemas-BuchanEmsian fauna were by Chapman (1916b), Hills (1936a, 1941), Woodward (1941) and White (1952). The Antarctic Aztec Devonian fish fauna, discovered during the British 'Terra Nova' expedition of 1910-13, was described by Woodward (1921). Hills (1929, 1931, 1932, 1936b) first recorded Late Devonian fish faunas (E Victoria and NSW). Teichert (1949) and Hills (1959) reported Devonian fish remains from NW Australia (Canning Basin) and central Australia respectively.

This early discovery phase was summarised by Hills (1958), who assessed the Early Devonian TaemasBuchan fauna ofSEAustralia as cosmopolitan at family level, and the Late Devonian assemblages as entirely cosmopolitan at generic level. He noted, however, that the association of three Late Devonian placoderm genera (Bothriolepis, Phyllolepis and Remigolepis) suggested that they did not have the same relative temporal significance in Australia as in Europe (subsequent1yconfinned by Young, 1974).At that time new discoveries in areas including Australia were expected to confirm and extend the knowledge of


Devonian fish faunas derived from scientific investigations in the Northern Hemisphere. This was overturned with Ritchie's (1969, 1973) investigations of the fish fauna from the Mulga Downs Group of\V NS\V, which revealed a fauna dominated by new taxa, including the endemic placodenn Wuttagoonaspis (Fig. 13J), of assumed Early-Middle Devonian age. Previously (Rade, 1964), a Late Devonian age was based on the assumed presence of a typical 'Old Red Sandstone' assemblage ofthe placoderms Phyllolepis, Holonema and GlVenlandaspis.

A more reliable taxonomic and biostratigraphic database for East Gondwana, applicable to questions of mid-Palaeozoic global biogeography, is now available (Long, 1982, 1991, 1993, 1995b; Tumer, 1991, 1993;Young, 1993a-b, 1995c;Youngefal., 1993; De Pomeroy, 1995, 1996; Turner ef al., 2000; Long & Trinajstic, 2000; Young & Turner, 2000). Collections in Australian instihltions contain many new taxa; discoveries continue to be made. The increase in taxonomic data fromAustralasia is part of a remarkable increase in knowledge of Palaeozoic vertebrates over the last 30 years, impacting significantly on global understanding of the biogeography of Devonian faunas (Young, 1990a, 1990b; Janvier, 1996).

Non-marine biota 15Tasch (1987) has reported the horizon ofCyzicus

talenti Tasch as being "Fairy Bed, Snowy Volcanics, Buchan Caves Limestone". The material was from dark mudstones in the Fairy Sandstone about 200 m upslope on the S flank of Spring Creek W of Buchan. This horizon is 120 m or more below the base of the Buchan Caves Limestone, with the strikingly red McRae's Ignimbrite (Orth ef al., 1995), infonnally known as the "salmon porphyry" by earlier workers, lying between the two. As the base of the Buchan Caves Limestone is believed to be within the pireneae Zone of the late Pragian (see above), the Fairy Sandstone is inferred to be Pragian, perhaps mid-Pragian.

Macrollor. (J.G. Douglas) 16Plant remains reminiscent of certain Northern

Hemisphere Carboniferous forms were noted from the Melbourne Terrane over a cenhtry ago (Nicholas, 1875), and referred to Lepidadelld1'01l. Vincent (1925) discussed a 'Psilophyton' flora from the Walhalla and Woods Point districts but did not clearly differentiate this assemblage ii-om Late Devonian floras described earlier from molasse sequences in Victoria and elsewhere in EAustralia. She discussed, inter alia, the key plant of the assemblage, 'Arthl'ostigma'. Other investigators, (see Harris & Thomas, 1941) had noted occurrences in several areas of a large leafY plant fossil associated with graptolites identified as being of late Silurian (Ludlow) age. The mystery 'Lepidodendron', 'Arthl'Ostigma', and "large leary plant fossil" was later identified as Baragwanathia longijolia by Lang & Cookson (1935).


In earlier papers (Lang & Cookson, 1927, 1930; Cookson, 1935). associated species were described, but the essence of this work was the presentation of a large land plant with well defined stem and leaf units and vascular tissue in SEAustralia at a time when elsewhere the land flora appeared to have been confined to very small stems bearing primitive sporangial sacs (e.g., Cooksonia, Lang, 1937). Baragwanathia became entrenched in palaeontological literature as a salient innovation in the histOlY of life.

The appearance of Baragwanathia naturally aroused interest in its ancestral line. Intense search in older strata has failed to produce any trace of immediate progenitors. Stems (Douglas, 1965) from the Serra Sandstone, Grampians Group, regarded as no younger than Early Silurian (Cayley & Taylor, 1997), though this seems debateable from radiometric data (cf. Mawson & Talent, 2000), may have affinities with undescribed zosterophyll components of the Baragwanathia flora.

The situation whereby an advanced vascular plant was present in such ancient (Ludlow) sediments, was made more acceptable to some by re-identification (Jaeger, 1966) of the prominent graptolite associate, 'Monograptus ullcinatus', as M. thomasi, a now well known Pragian fonn (Jaeger, 1966). This dating, was further strengthened by description of Pragian conodonts from the Wilson Creek Shale, a unit with M thomasi in the Tyers River area (Carey & Bolger, 1995).

17Baragwanathia has been depicted (Douglas, 1983), as a pioneer colonizater of the land in estuarine situations, whereas others have proposed fossilization tens or even hundreds of kilometres offshore after being swept out to sea by large rivers.

18Considerable new information on the plants in the younger (Devonian) part of the sequence has been obtained from numerous other localities in the YeaAlexandra-Eildon area (Couper, 1965). The new localities include Yelland Track, stratigraphically close to the topotype 19 mile Quarry locality.

Tims (1980) described the anatomy of Barag'Nanathia stems from Boola Quarry (pragian) with "much better preservation than the specimen from which Lang and Cookson derived the anatomy", and confimled allocation to the Lycophytina.

19Douglas (1983) in a pictorial reconstruction presented a scene not unlike that envisaged for the much better known forests of the Northern Hemisphere Carboniferous, with robust woody Leptophloeum alfstraie McCoy dominating the scene.

251

Charophyta (M. Feist) 19Entry of Icriodus symmetriclfs has been taken to

indicate Frasnian as opposed to Givetian, but caution is necessary because of a very few, though undocumented, late Givetian reports of this species (Mawson & Talent, 1997). An erosional surface between the top of the Stanley Limestone Member and the base of the redefined Mytton Formation implies a hiatus between the two units, at least locally (Feist et al., 2000), but what it means in terms of time is problematic.

,

I , I ,

Rockhampton I , Queensland

I 30 ,

-1_--l , Brisbane

33

o

I 28 ~------f-1';--I New South Wales 27

I 21 16 19

'L 22 "I , 1; 7'~' I '.r\ 15· •

, Vic. "'""'--, '12 I 4 ,5 -, • ..R.6 • _',>'<"'e~

(!\f.4l..... 7 MelbOU{ 8

~ ~T.,m'ni' 500km 1VCObart

t I

26 25 24 23 20

Location of the main eastern Australian Devonian sequences (from Mawson & Talent, 2000). Commentary on the sequences shown here is given by Mawson & Talent (2000).

252

®

\

1"""" r '..r--. Wagga , Waggaj

'I "-.~. {'. -~ t; •••••

'(II ;.~ .• ~ ~/Buc~an ..... I ~ -;j("fi". "U·, ...

Melb0lJ!9-e~ {l: :~ <'?66erabbera :\; "l ' .. '_...." \'~ ?i ., ...... , .... ;, ~

Or ; II .0 10. "' ... I'---. .. 0 -'

I \.)~~ ~ ... ~ "'~ Queenslown\"{oUA'~ ~ ~ ~

\\, " ~ ,?-... .;t, Hobart ~ <a. .. ~

9:. ..


® \'

\\~ l~ Calms Chillagoet1

1-.. ...... ':::::;-• ., : •••• ~ownSvil!e ;"., .. , Ii

Charters· ~~ Towers I~\

\t Emerald. ". Rockhampton

"<;" 1. __ . Monlo. \.\

---1 Adavale. ....

l Roma'!I , t:I.:' ) Goondiwindi ,_Ii ./ ~ r·-·--~:r:. It i Cob~ ~ If Tamworth

\

1 Ir· .. ·····~ney r'..r- Wagga \: f 8Yd

, ',,-Wagga' i 0( c7~ nberra

1 '~'/.i I '.~ , "J ,. , I ".i_Suchan

MelbOU[ge~,<~;a~era

t N

I

° '-/ll"0%~ QUeenslow\1. \'''~ ~.;.C> .. ~ ... ~

~. ~~ Hobart ~'"

"'~ o~

'" o ~ ~~~,.;5;;;OOkm

Extent of marine transgressions for selected intervals of Early Devonian time in E Australia. Note postulated extension of marine conditions as far W as the Darling Basin) based on wide distribution of sulcatus Zone and possibly pesavis Zone through western NS\V.

AAPMemoir23 (2000)

,

I I I i

®

Melbourne p ~

QUeen"ow~ ,nUbart

,

I , I ,

® I

\1

\\ Chmagoe. U

" : ,. . ........... , ~.~ Towjlsville -Charters·

Towers

I , Emerald, \. • Aockhamplon

I " Mon}~. , -.l·-·l Ad".'e, ,1 I

Roma-Brisbane' l ~ I Goondfwlndi ~ I _

~·-·-·--../1',.I" I Moree' j,

Gobar •

\' '\<' '2::: r'.;-- Wagga :. ;;'SYd I ,Wagga· .~ I "'-_..->.._ ?anbe~a

! Tabbarabbe;l~ 1 \ I '. .Buch~n Me[bo\!!!>e~ \ ~

t N

"";r- 2t p ~ \\~ ~

QUeensl0V." , \\ \ lm -:;. Hobart -:. ~

~ .. ~ ~

253

~ -:a.

I ;;.0 ~~~~5;.;.00.km

~

Extent of marine transgressions for selected intervals of Middle and Latc Devonian time in E Australia. Note postulated extension of marine conditions into the Adavale Basin during the Eifelian, and the almost balTed nature of the Burdekin Basin (inland from Townsville) during the early Givetian.

ai > . sandt;;;Oi

,. 12• Zeehan

Point Hibbs - .. - - .

3. North East Tasmania

4. The

Grampians

5. HeathcoteRedcastle Lilydale

17. 1

8.

_ Wa,I,hal!a WaS"'tah ay

1 9 I'~. • Mitch~.11 R. Bucha~ _ .. 1 R. Bindl

6.

[C ~ ~ U. du JICl1W : « ~ {!. Ldupllco.ta :, i ,

11. Boulder

Flat

o sufcattJ """, I

I Praosufcata - - -+- - -:--- -4-- - +--- -:---+ ---+----:- --+ ---

, I I , , , I I I

oxpans~ ::::::::: :: :::', ... : .... ' I "" ?~? . ?---L...?....l , " r .---y-. I

mum/nlfors. om O! DlI

croplda W trlnn u/ar/s I- In U {ormis ::5 rhenana

Avon R. Gp

:ii omlOaQ '2 I I,', ',: 't' ~uncata ::::::

Z u. tmm;/tnn, ::::::

MtTambo Gp Gp'

?

« fa/slovalls :::::" _ ~__ ___ _ __ ~ __ ~ ___ ~ __ ~___ _ __ ~ __ ""; ___ L __ _

- e dlsarflfs ' , , <II ' , ,

Z _ fJrmflnni ' , ,

W <II ' , , .,J.2: V!lfCUS ' , I

o 0 (:I hem{ansalus \ \ \ > ~ onsons/s s.s. Tabbe'rabberan Events : UJ ::s .§ kockellanus : : :

] australis : : : Q w cosl8tus ' I ,

(lft/tus I I ,

<lUllS -- --- ---:-----t---t----i---" ---t----,--- .... ---~ sorotlnus :: :Cavos Hili Ss. : : Murrlndal ~ ?---i .- InvefSus-laticost. Whltohorso , , bL ~?? ,Ls Taravalo Fm E BoaehSs.'" I";":' , w perbonus-gron. and Rod Roof? ?:: Cathedral G Uptrap Fm ?-?

~ dehlsoons Cliff Sit : :' , , Walhalla Go Wentworth i D! IronOQO f f' Roeklands ' i-?--? if ~ . Gp <C <0 kindlei Rh lit I Ulydale Ls. ~ (I) ~ Boll Pt Ls. lot W c: sulcatus ptHibbsFm. r yo III ?_? ~i3 &13 WaratahLs \

:> OSBVlS 't -w -----.:' ? __ ?- . ? . WlI~Horso

eurekaensls Grampians Mt Ida fm. Fm.

Snowy A Voles

.......

. -7-'[Bouldor F1a! Ls.J

[Bungywarr Fm.]

n delta : ? ' '?1' Humevale Boola Bods ,m

...I woschmldtl Bell Shale Mathinna Gp ?? Whitelaw Stt. - - Beds .--. ,

Prld oostolnhomonsls ~~~c;:, r?? Mcivor F~.- - - - -I +7 ? .. ~ - - -: - --~ ....... 9 t:vents -i---;----

.J w ~ crIs us I-?? . ~. ,? .?- t t : CIJ !;;: ~ Ia.tlalata: : T Sinclair : I Enano

...J':: sliurlcus, ,Melbourne Valley SIt. : Gp ooonsis' : Fm. , I I

Devonian stratigraphic columns from Tasmania and Victoria. Numbers (1-11) refer to localities on the map of eastern Australia at the beginning of this appendix.

N V> ...

~ f =;.

~ D §

12. 18.

Ravine Limekilns

ai > . , a:: ~ ~ U.du Ilcat8 :

I ~ <[ {2 L dupllcata : OW sulcatlJ ' , I , , , ,

--~---~---+---7---7---7---~---I praesulcatB , , I , I , ,

, I , I , I I _u_____ :: :

Z

<t

Z

0

> w

C

W

~ ...

W

-' 0 0

"

~ II:

'" W

...ilw Ci.i !;;: ...

, " , " " " , ? __ ?'? ? I ,

[Men!l'pbUlaIcatombal Gp . ] i

, ?--? ? ? , ' , ,

, : : : ' , I

-------~---~---~---~---+---~---

-E=!'-'"' ~ I Hatchery Ck afiljUS r Fm.

I "~:;;~:: Aooo' Too Fm ;'e:~s ~s~

Milk Shanty Fm

volcanics volcaniclastics

ao' clastics

: : : : : , " , " , " : '

,

Tabberabberan Events , : ) diachronisim I ___ _

-~~~~~~~~~~~~--~y-,

Majurgong : Fm.

Tangerahg Fm.

Mandagery Pk Fm.

Goonigal Gp

Red Hill Mbr

Cunningham Fm

20. Cudgegong

RyJstone

Devonian stratigraphic columns from New South Wales. Numbers (12-20) refer to localities on the map of eastern Australia at the beginning of this appendix .

~ ~ ~. '" w

)

N

'" '"

ai a: ~

Z

...

~ II:

Ul

w l;: ....

Zlw

o > w Q

.... c c ::;

~

~

.Jlw (i) ~ ....

Pigna

, , , , , ",

---~-----+---+---+---~-----

' "" : :::: ~I : t? ?t i ' i ! 3f'

Hervey Gp

,

, , ,

Mostyn Vale Fm.

---.,.--------------

Mulga Downs Group

, :

Dulladeny Voles

?

Glen Ward Beds

Pitch Ck Voles

Timor Ls. Mbr

Yarrimie Formation

, , , ,

'GJencairn : Ls. :

28. Sliverwood,

Etc.

Keepit Congo

Kopyje, Ootha & Derriwong Groups " ,

...__._?-----r- - - - -:- - - - +- - - - - - - - -+ - --" , " , " , " , , , , ,

Devonian stratigraphic columns from New South Wales and Queeusland. Numbers (21-28) refer to localities on the map of eastern Australia at the beginning ofthls appendix.

'" V>

'"

~

f. tv W

'i0 <::> <::> .s

ai a: (l

~ 0: .. W

~ 0: .. W

-'IW U5 ~ ~

Telemon Fm.

,

: : : : ---r--~---~---~---

" ' " ' " ' " ' " ' " ' " ' :: i-L-?~ '? ?:?? IGr~y~ankl

Bundock Ck Gp

Fm.

Vanneck, Stud and

Julia Formations

c

~ E

"' ~ " -g :c

.......-:: +--. Big Bend f---=--"'ij Arkose

,

, , ,

Gumbardo Fm. I Beds

~?- - ~?-

, T , , Shield Ck. Fm. --f;:_~ _ I

> ' , D3--~ delUl : Martins Well t? Ralph Flint: '£ flufekoensls I : , , and Arch Ck Mbr : .3 woschmldtl : : : : _ Is. mbrs erosional interval :

----~--' , , " , Prld oostelnhomensls ' -..- - - - r-- - - -,- - - -,- - - -r---=-=,I..- - - -'- --

, I , " " , ' , I' ' , ' , , , , , I , , , , , Jack Fm.

: I ,

~ F :2 "

Devonian stratigraphic columns from Queensland. Numbers (29-38) refer to localities on the map of eastern Australia at the beginning of this appendix.

~ ~ ~. t:l

~ -9

tv ~

"

Devonian palaeobiogeography of Australia and adjoining regions

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