Middle Palaeozoic microvertebrate assemblages and biogeography ...

Abstract 

Available online at www.sciencedirect.com 

Palaeoworld 19 (2010) 37–54 

Research paper 

Middle Palaeozoic microvertebrate assemblages and biogeography 

of East Gondwana (Australasia, Antarctica) 

Carole J. Burrow a,∗ , Susan Turner a,b , Gavin C. Young c 

a Geosciences, Queensland Museum, 122 Gerler Rd, Hendra, Brisbane, Queensland 4011, Australia 

b School of Geosciences, Monash University, Victoria 3800, Australia 

c Research School of Earth Sciences, The Australian National University, Canberra, Australian Capital Territory 0200, Australia 

Received 22 December 2008; received in revised form 18 October 2009; accepted 1 November 2009 

Available online 11 November 2009 

Silurian vertebrate remains are rare in the Australasian region, mostly lacking from the end Ordovician to the mid-Ludlow presumably because 

of the purported Gondwana Ice Age. Thelodont, placoderm, acanthodian, ?stem actinopterygian, and probable chondrichthyan remains are known 

from eastern and western Australia and Irian Jaya. Of significance are the links between eastern Australia and China, with sinacanthid spines 

known only in these regions, and both having porosiform, but not punctatiform, poracanthodid acanthodians, while Western Australia, Iran and 

possibly Irian Jaya have similar thelodonts, all from shallow marine to evaporitic settings. Earliest Devonian (Lochkovian) vertebrate microfossils 

include placoderm taxa with circumArctic and Bohemian affinities, but post-Lochkovian marine assemblages comprise mainly endemic forms of 

turiniid thelodonts, placoderms, acanthodians, chondrichthyans, and sarcopterygians. Most of the Early Devonian marine assemblages from eastern 

Australia indicate tropical–subtropical depositional environments. From the Middle–Late Devonian, notable index taxa include Phoebodus spp. 

After the Frasnian-Famennian events, turiniids finally disappear in the Australian record, placoderms also become absent by the end of the period, 

acanthodians are increasingly dominated by acanthodiforms, and chondrichthyan and actinopterygian diversity increases. For the Carboniferous, 

vertebrate occurrences are early to mid-Mississippian, disappearing in the early Pennsylvanian in marine and non-marine environments. 

During the time span of IGCP 491, data on previously poorly known and many new taxa represented only by isolated remains have been 

analysed, and a wealth of new acanthodian taxa, a new Early Carboniferous tetrapod Ossinodus pueri, and several sarcopterygian taxa have been 

fully described. 

© 2009 Elsevier Ltd and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved. 

Keywords: East Gondwana; Palaeobiogeography; Palaeobiostratigraphy; Palaeozoic; Vertebrate micro-remains 

1. Introduction 

The broad picture of the vertebrate fossil record for the 

Middle Palaeozoic of East Gondwana shows the paucity of 

Australasian Silurian assemblages (Burrow and Turner, 2000; 

Burrow and Turner in Pickett et al., 2000; Burrow, 2003a), followed 

by an endemic radiation of Devonian thelodont agnathans 

and a dramatic increase in gnathostome (jawed vertebrate) diversity 

with the Devonian ‘Age of Fishes’ radiation. After the rise of 

tetrapods by the beginning of the Late Devonian and the demise 

∗ Corresponding author. Tel.: +61 7 33916626; fax: +61 7 3846 1918. 

E-mail addresses: carole.burrow@gmail.com (C.J. Burrow), 

paleodeadfish@yahoo.com (S. Turner), gavin.young@anu.edu.au 

(G.C. Young). 

of the placoderms before the Devonian-Carboniferous boundary, 

both marine and non-marine Carboniferous assemblages are 

characterized by significant radiations of chondrichthyans (cartilaginous 

fishes) and actinopterygians (ray-finned bony fishes). 

These occurrences are mainly restricted to the Mississippian 

(Turner in Jones et al., 2000a; Turner et al., 2008), before the 

onset of cooler temperate to cold conditions towards the end of 

the Viséan, leading up to the Permian glaciation of the southern 

continents. 

Significantly absent from the Australasian Middle Palaeozoic 

are various groups of armoured agnathans, a major component of 

Silurian and Devonian fish faunas in the Northern Hemisphere 

(e.g., Blieck et al., 2000). Of agnathans, apparently only the 

thelodontid and turiniid thelodonts and the enigmatic pituriaspids 

occur in the Siluro-Devonian of Australasia, with only 

the thelodonts preserved as micro-remains, viz. scales (e.g., 

1871-174X/$ – see front matter © 2009 Elsevier Ltd and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved. 

doi:10.1016/j.palwor.2009.11.001

38 C.J. Burrow et al. / Palaeoworld 19 (2010) 37–54 

Turner and Burrow, 1999). However, the gnathostomes are abundant 

and diverse, dominated by placoderms in the Devonian but 

with many osteichthyans, acanthodians, and chondrichthyans (in 

order of decreasing known diversity). Vertebrate micro-remains 

are common, in both geographical and stratigraphic distribution 

(e.g., Turner et al., 2000; Young and Turner, 2000; Basden et al., 

2000, 2006; Burrow, 2002, 2003b) and have been successfully 

used for biostratigraphical correlation schemes (e.g., Turner and 

Burrow, 2000). 

For the Carboniferous, vertebrate occurrences are mainly of 

early to mid-Mississippian age and include the only known pre- 

Permian tetrapod-bearing assemblage in Australasia/Gondwana 

(e.g., Warren and Turner, 2004; Warren, 2007), with vertebrates 

disappearing, however, in the early Pennsylvanian with a last 

record in Queensland (Turner in Jones et al., 2000a). Marine 

assemblages from northwestern and northeastern Australia are 

dominated by shark and actinopterygian micro-remains (e.g., 

Chambers, 2002; Trinajstic and George, 2005, 2007), and nonmarine 

assemblages in Victoria and central Queensland contain 

several sharks, gyracanthid and other acanthodians, rhizodonts, 

lungfish, and actinopterygians (Turner et al., 1999, 2005; Turner 

in Jones et al., 2000a; Johanson et al., 2000; Kemp, 2002; Parker 

et al., 2005; Garvey and Turner, 2006). 

Previous summaries of the biogeography of Middle Palaeozoic 

vertebrates were part of a review covering the total 

Phanerozoic fossil record for Australasia (Wright et al., 2000), 

including contributions on the Silurian by Pickett et al. 

(2000), the Devonian by Talent et al. (2000), and the Carboniferous 

by Jones et al. (2000a). Particular aspects of the 

overall composition, biogeography, and stratigraphy of Middle 

Palaeozoic vertebrates of Australia, with the emphasis on 

the record of Devonian vertebrate micro-remains, were covered 

in chapters of Blieck and Turner (2000), the final volume for 

IGCP 328: Palaeozoic Vertebrate Biochronology and Global 

Marine/Non-Marine Correlation, and by Basden et al. (2006). 

The Australasian fossil record for the Silurian and Carboniferous 

is much less prominent than for the same time interval in other 

areas. By far the greatest Palaeozoic diversity of both microvertebrate 

and macrovertebrate fossil assemblages preserved in East 

Gondwana are from the Devonian; Devonian macrovertebrates 

are covered separately by Young et al. (2010). The present paper 

deals with updates on the Silurian to Carboniferous vertebrate 

records, and a summary of recent work on Devonian microvertebrates. 

Institutional abbreviations: AMF, Australian Museum Fossil 

collection; ANU V, ANU College of Science Palaeontological 

Collection, DA Brown Building; GSWA F, Geological 

Survey of Western Australia collection; ML, Mike Leu collection, 

Sydney; MMMC, Geological Survey of New South Wales 

(GSNSW) collection; NMV P, Museum Victoria Palaeontology 

collection; RGM, National Natuurhistorisch Museum, Leiden, 

Netherlands; UQY, University of Queensland Geology collection 

(reposited in QM; GWK, Greg Webb subcollection); QMF, 

Queensland Museum Fossil collection. 

Fig. 1. Australian fossil vertebrate localities of Silurian and Carboniferous age referred to in the text. State abbreviations are: N.S.W., New South Wales; N.T., 

Northern Territory; Qld, Queensland; S.A., South Australia; Vic., Victoria; W.A. Western Australia; Tas., Tasmania. Geological province and sedimentary basin 

abbreviations are: AB, Amadeus Basin; ADB, Adavale Basin; BPB, Bonaparte Basin; BrR, Broken River Province; CB, Canning Basin; CAB, Carnarvon Basin; DB, 

Darling Basin; DrB, Drummond Basin; GB, Georgina Basin; LFB, Lachlan Fold Belt; OB, Officer Basin; THB, Timbury Hills Basin. Silurian localities (squares), 

numbered as in Pickett et al. (2000, fig. 4) are: S1, Bullock Creek, Broken River area (Qld); S2, near Cumnock (NSW); S3, Cheeseman’s Creek, near Orange (NSW); 

S4, east of Trundle (NSW); S5, Tharwa, near Canberra; S6, Cowombat area (Vic.); S7, Yea (Vic.); S8, Grampians (Vic.); S9, Kempfield 1 well, Canning Basin (W.A.); 

S10, Pendock 1A well, Carnarvon Basin (W.A.); S11, Yaringa 1 well, Carnarvon Basin (W.A.). Carboniferous localities (triangles) are numbered from the southeast 

in an anticlockwise direction as follows: C1, Mansfield (Vic.); C2, New England area (NSW); C3, Murgon (Qld); C4, Rockhampton area (Qld); C5, Narrien Range 

and Ducabrook areas, Drummond Basin (Qld); C6, Bonaparte Basin (W.A.); C7, northern Canning Basin (W.A.); C8, Broken River region (Qld).

2. Silurian vertebrates (see Fig. 1 for localities) 

2.1. Palaeolatitude 

The continental reconstructions of Scotese and McKerrow 

(1990) and Li and Powell (2001, fig. 11) placed Australasia in 

the tropics, entirely between 20 ◦ N and 20 ◦ S throughout the Silurian, 

on the eastern side of Gondwana. New Guinea (with Irian 

Jaya) was located nearby and New Zealand was much closer to 

Australia. Carbonate platforms fringed the eastern coast, along 

the western margin of the Tasman Fold Belt, but the main deposition 

was siliciclastic. Shallow-marine carbonate and evaporite 

deposition occurred in the Canning and Bonaparte Basins of 

Western Australia; central Australia was probably also arid. The 

North and South China Blocks were placed at a similar latitude, 

scattered to the west of Australia. The closest depositional zones 

to the east were in the Palaeo-Pacific oceanic basins (now totally 

disappeared), and in the present-day Nevada region. 

2.2. Vertebrate records 

All known surface occurrences in Australia are in the Tasman 

Fold Belt, with the only other records being from boreholes in 

Western Australia. The one record of a limited assemblage from 

Irian Jaya includes thelodont scales; those originally referred 

by Turner et al. (1995) to Thelodus trilobatus might be better 

placed in Praetrilogania (Märss et al., 2007). As yet there are 

no known remains in New Zealand. 

The major groups represented are the Thelodonti and Acanthodii, 

with very rare occurrences of putative stem osteichthyans: 

Lophosteus from the Ludlow Jack Formation in the Broken River 

region of north Queensland, and possibly Andreolepis or Ligulalepis 

(Fig. 2A) from the Pendock-1A borehole in Western 

Australia (the assemblage from the latter awaits full systematic 

description). The acanthodian from this sample is tentatively 

assigned to Nostolepis cf. alta (Fig. 2B), and the few thelodont 

scales resemble some figured by Hairapetian et al. (2008, fig. 

4D) from the mid-Silurian Niur Formation, Derenjal Mountains, 

Iran, which they assigned to Loganellia sp. cf. L. grossi (Fig. 2C 

and D). One fin spine from Irian Jaya (Turner et al., 1995, fig. 6) 

might belong to a putative chondrichthyan, perhaps a primitive 

holocephalan, by comparison with the surface textures in Mesozoic 

spines (ST, pers. obs.); associated vertebrate micro-remains 

include thelodont (Fig. 2P) and acanthodian scales (Fig. 2Q). 

Few of the genera found are endemic, with most also occurring 

in the present circumArctic region, e.g., Nostolepis and 

ischnacanthiform acanthodians (Fig. 2B, E–J, N, O and Q), and 

probable turiniid (Fig. 2K and P) and thelodontid (Fig. 2R–U) 

thelodonts. The assemblage from the?late Ludlow Silverband 

Formation, Victoria is dominated by the endemic thelodont 

‘Turinia’ fuscina Turner, 1986 (Fig. 2K). The generic identification 

of this species requires reconsideration, but the material 

is not amenable to histological study being preserved mostly as 

natural casts, or friable white scales in sandstone. Body scales 

referred to a new taxon Niurolepis susanae from the Niur Formation 

(Hairapetian et al., 2008, fig. 2) share similarity with 

‘T.’ fuscina; associated gnathostome elements (Sinacanthus?fin 

C.J. Burrow et al. / Palaeoworld 19 (2010) 37–54 39 

spines and Radioporacanthodes scales and tooth whorls) show 

a Chinese affinity (Turner, 1986; Burrow, 2003a; Fig. 2H–J). 

The largest known fossil is the articulated pectoral and partial 

trunk region of a large teleostome fish, Yealepis douglasi 

Burrow et Young, 1999 (Burrow and Young, 1999, Fig. 2L 

and M), which lacks any evidence for fin spines but has a 

typical narrow-based acanthodian pectoral fin and squamation. 

Scales of the same form as on Yealepis are found in microvertebrate 

assemblages from the Lochkovian-Pragian of eastern 

Australia (Fig. 5H), the Pridoli-Lochkovian boundary beds of 

the Birch Creek Section (Roberts Formation) in Nevada, USA 

and the Klonk Section (including the GSSP for the Silurian- 

Devonian Boundary) in the Czech Republic (CJB, pers. obs.), 

the Lochkovian of arctic Canada (Vieth, 1980), Taimyr, Russia 

(Valiukevičius, 1994), and the East Baltic and Byelorussia 

(Valiukevičius, 1998), as well as the Pragian-Emsian of China 

(Burrow et al., 2000). Scales from the circumArctic localities are 

invariably small compared with those from Australia, Nevada, 

the Czech Republic, and China, and are assigned to Nostovicina 

laticristata (Valiukevičius, 1994). However, the type species 

for Nostovicina Valiukevičius et Burrow, 2005 (Valiukevičius 

and Burrow, 2005), a genus based mainly on histological structure 

of scales, is N. fragila (Valiukevičius, 2003), which has 

fin spines. The Lochkovian MOTH locality in the Northwest 

Territories of Canada has yielded one articulated specimen comprising 

the head and branchial region with N. laticristata scales 

and which, like Yealepis, lacks fin spines (Hanke, pers. comm., 

2000). Other circumstantial evidence (e.g., micro-remains which 

include this form of scale plus poracanthodid scales, but only 

poracanthodid fin spine fragments) indicates that Yealepis and 

a subgroup of “Nostovicina” comprise a group of acanthodians 

that lost fin spines, or less likely, a group of teleostomes 

that never had fin spines. The latter possibility seems least 

likely because fin spines or spinal plates have been identified 

in all other stem groups of gnathostomes. Morphology, average 

size, and histology all indicate that the Czech, Nevadan, 

and Australian scales are conspecific, supporting evidence from 

invertebrate fossil groups (cf. Pickett et al., 2000), which suggest 

faunal interchange between these regions during the Late 

Silurian-Early Devonian. Another scale-based poracanthodid 

acanthodian genus Trundlelepis (Fig. 5I) is found in the Silurian- 

Devonian boundary beds in Nevada (CFB, pers. obs.) as well as 

in the?late Lochkovian of southeastern Australia. 

Possibly the youngest Silurian assemblage, comprising probable 

thelodontid thelodonts (Fig. 2R–U) and indeterminate 

vertebrate dermal bone and possible acanthodian fin spine 

(Fig. 2V) fragments, is from the Kempfield 1 bore hole 

(?Pridoli), “Carribuddy” Formation, Western Australia (Turner, 

1993a). The Upper Silurian-Devonian(?) Caribuddy evaporites 

formed a thin stratum with restricted basin development within 

the northern Fitzroy Trough and Kidson Sub-basins (Reilly, 

1988). The specimens are few and extremely abraded, mostly 

precluding more precise systematic determination; the thelodont 

scales, however, are best preserved and might represent a new 

species, reminiscent of those of the type species Thelodus parvidens 

and even the head scales of Niurolepis susanae of Iran 

(Hairapetian et al., 2008), or some of those described as species


Fig. 2. Silurian vertebrates from Australia. (A–D) Scales from locality S10. (A) ANU V3460, early actinopterygian Andreolepis or Ligulalepis sp. caudal scale, 

crown view; (B) ANU V3461, acanthodian Nostolepis cf. alta scale, anterior view; (C) thelodont? ANU V3462, crown view; (D) thelodont? ANU V3463, lateral 

view of detached crown. (E and F) Gomphonchus? turnerae holotype scale UQY7707 in crown view, and tooth whorl UQY7697 in occlusal view, from the Jack Fm.,

Fig. 3. Late Silurian-Early Devonian distribution of poracanthodid acanthodians 

and Silurian Gondwanan turiniid and thelodontid thelodonts, plotted on 

the global biogeographical reconstruction of Li and Powell (2001, fig. 12) for 

the Early Devonian (400 Ma). (♦) Silurian Gondwanan thelodonts; (○) Silurian 

porosiform poracanthodids; (△) Devonian porosiform poracanthodids; (�) 

Silurian/Devonian punctatiform poracanthodids; (*) Silurian/earliest Devonian 

machaeracanthids. 

of Parathelodus from the Late Silurian of China (Märss et al., 

2007). 

2.3. Palaeobiogeographic significance (Silurian) 

Pickett et al. (2000, figs. 1–3) noted that Australian rugose 

coral assemblages show a ‘double affinity’, with western and 

arctic North America assemblages on one hand, and Asia on 

the other. Australian vertebrates show a similar duality, with 

the western USA-circumArctic-Bohemian affinity noted earlier, 

and also a Chinese affinity. In addition, there is new data 

from across North Gondwana (e.g., Iran as noted above, and 

older Ordovician vertebrates from Oman: Sansom et al., 2009), 

which requires assessment. Sinacanthus spines are recorded 

from many local Chinese basins (Zhu, 1998; Sansom et al., 

2005), but otherwise only (if somewhat tentatively) from Australia 

(Burrow, 2003a). Distinctive scales from poracanthodid 

acanthodians show a wide geographical range in the Late Silurian 

to Early Devonian (Fig. 3). Whereas both punctatiform and 

porosiform poracanthodid acanthodians are found in Laurussia, 

only porosiform poracanthodids have been found in Australasia 

and China, from the Ludlow through to the Emsian. The only 

Australian Silurian poracanthodid identified (Parkes in Basden 

et al., 2000; Burrow, 2003a; Fig. 2J, N and O) is very close to 


Radioporacanthodes qujingensis (Wang and Dong, 1989) from 

China. Burrow and Young (2005) suggested that Machaeracanthus 

Newberry, 1857 (Newberry, 1857) could have derived 

from a poracanthodid; earliest occurrences of Machaeracanthus 

scales are in the Siluro-Devonian boundary beds at Klonk, Czech 

Republic (CJB, pers. obs.) and in the Pridoli of eastern Canada 

(Legault, 1968). 

We have noted above the similarities becoming apparent 

between Ludlow-Pridoli thelodont scales in Western Australia, 

Iran and South China (Fig. 3). The limited Irian Jaya 

record, however, shows affinity with Baltic-circumArctic assemblages; 

more data from the New Guinea-Indonesian archipelago 

would help to define the palaeobiogeographic relationships. The 

present evidence supports a climatic, palaeolatitudinal distribution 

along and off the northern margin of Gondwana, possibly 

due to longitudinal marine palaeocurrents. 

3. Devonian microvertebrate assemblages (see Fig. 4 for 

localities) 


Limestones in Australasia, including some with welldeveloped 

coral reefs, occur in the Early Devonian (e.g., 

Burrinjuck, NSW, Buchan, Victoria, Reefton, New Zealand), 

Middle Devonian (e.g., Broken River, Queensland), and Late 

Devonian (e.g., Canning and Bonaparte Basins, northwestern 

Australia). Only one significant New Zealand record is known 

from the Reefton Beds limestone (Macadie, 2002). All these 

generally indicate low paleolatitude tropical–subtropical climates. 

Li and Powell (2001, fig. 12) placed Australia between 

10 ◦ S and 40 ◦ S, on the eastern margin of Gondwana and relatively 

widely separated from the North and South China 

Blocks, which were both above the equator at similar palaeolatitudes. 

The increase in occurrence of presumed freshwater 

(or very shallow water marine) assemblages in Australia, and 

their appearance also in the more southerly Antarctic portion of 

East Gondwana during the Middle Devonian provides evidence 

of higher rainfall compared with a presumed more arid climate 

during the Early Devonian (Young et al., 2010). 

3.2. Early-Middle Devonian 

3.2.1. Marine assemblages 

Thelodonts are generally represented by endemic turiniid 

taxa (e.g., Turner, 1997a; Turner et al., 2000). Placoderms 

locality S1. (G) Ischnacanthid,? Gomphonchus sandelensis fin spine and scales ANU V1637 from the Tharwa Shale, locality S5. (H–K) From the Silverband Fm., 

locality S8. (H and I) (both to same scale), Sinacanthus? micracanthus holotype NMV P14364 and paratype NMV P14365 fin spines; (J) Radioporacanthodes cf. 

qujingensis tooth whorl (in oblique natural section), adjacent to?pectoral fin spine on NMV P14363; K, thelodont ‘Turinia’ fuscina scale NMV P207257.7, crown 

view. (L and M) Yealepis douglasi partial articulated fish lacking head and tail ANU V2351, specimen part (edges of specimen marked by white line), SEM of cast of 

scale impressions on counterpart, from the Yea Fm., locality S7. (N and O) Radioporacanthodes cf. qujingensis scales AMF97980, AMF97981, crown views, from 

unnamed Mb. (Ludlow-Pridoli), Garra Lst., locality S2. (P) Thelodont ‘Turinia’ sp.? scale RGM384.621, anterocrown view, and (Q) RGM384.626, acanthodian G. 

sandelensis scale, crown view, from locality 46, Fig. 4. (R–V) Scales and fin spine fragment from locality S9. (R–U) Thelodontidid thelodont scales: (R) GSWA 

F51458, head scale in basal view; (S) GSWA F514598, head scale in laterocrown view; (T) GSWA F51460, head scale in laterobasal view; (U) GSWA F51461, 

head scale in lateral view; (V)?acanthodian fin spine fragment GSWA F51462, leading edge view. fs, fin spine; arrows point to anterior. Scale bar = 100 �min(A–E, 

N–Q); 200 �m in (F, R–U); 500 �m in (K and V); 1 mm in (G–J and M); 1 cm in (L); (H and I) equalized in Photoshop ® .


and acanthodians from the earliest Devonian (early Lochkovian) 

show a mix of cosmopolitan and endemic taxa (marine 

deposits of New South Wales; Burrow and Turner, 1998, 1999; 

Parkes in Basden et al., 2000; Burrow, 2002, 2003b, 2006). 

The acanthodians include scales of species first described 

from the circumArctic Laurussian region: Nostovicina lacrima, 

Gomphonchus sandelensis, Gomphonchoporus hoppei, Radioporacanthodes 

porosus, and Zemlyacanthus menneri (Fig. 5F). 

These taxa are also found in coeval deposits in Nevada, USA 

(CJB, pers. obs.). There are several forms of acanthothoracid 

placoderms, and early representatives of endemic placoderm 

taxa (Burrow, 2003b). The mid-Lochkovian is marked by the 

appearance of the endemic putative palaeoniscoid Terenolepis 

turnerae at the base of the Connemarra and Garra formations. 

Scales of porosiform poracanthodids including Radioporacanthodes 

(Fig. 5G) and Trundlelepis cervicostulata Burrow, 1997 

Fig. 4. Locality map for Devonian microvertebrate occurrences of Australasia, and Silurian/?Devonian of Irian Jaya, mentioned in the text. All Australian Devonian 

vertebrate localities numbered; those represented only by microvertebrate assemblages indicated by black triangles; associated micro- and macroassemblages 

indicated by black circles; just macro by black squares. An additional microvertebrate locality assigned to East Gondwana (not shown) is the Aztec fish assemblage 

of southern Victoria Land, Antarctica (see Young et al., 2010, fig. 1, locality 45). Abbreviations for countries and Australian states are: IJ, Irian Jaya; N.S.W., New 

South Wales; N.T., Northern Territory; NZ, New Zealand; PNG, Papua New Guinea; Qld, Queensland; S.A., South Australia; Vic., Victoria; W.A. Western Australia; 

Tas., Tasmania. Geological province and sedimentary basin abbreviations are: AB, Amadeus Basin; ADB, Adavale Basin (subsurface Devonian); BPB, Bonaparte 

Basin; CB, Canning Basin; CAB, Carnarvon Basin; BT, Bancannia Trough; DB, Darling Basin; GB, Georgina Basin; LFB, Lachlan Fold Belt; OB, Officer Basin; 

THB, Timbury Hills Basin (subsurface Devonian). Locality details (numerical order) are: 1, Mount Howitt/Tatong; 2, Buchan/Bindi, Gippsland; 3, Yea area; 4, 

Eden/Pambula area; 5, Taemas/Wee Jasper/Tumut/Wellington area; 6, Forbes/Jemalong area; 7, Canowindra; 8, Gap creek, Orange; 9, Grenfell; 10, Troffs/Trundle; 

Bogan Gate; 11, Keepit Dam; 12, Gunderbooka/Cobar; 13, Wuttagoona/Tambua/Mt. Jack; 41, Broken Hill area (Barrier Range, Mutawintji); 14, Adavale subsurface; 

15, Mt. Beaufort, Drummond Basin; 16, Mt. Podge and Fanning River, Burdekin Basin; 17, Broken River area; 18, Gilberton, Georgetown Inlier; 19, Chillagoe area; 

20, Toomba Range/Cravens Peak; 21, Dulcie Range; 22, Ross River Syncline; 23, Stokes Pass; 24, Mount Winter; 25, Bonaparte Basin; 26, Wilsons Cliffs 1 well; 

27, Gogo-Lawford Range area; 28, Mt. Percy; 29, May River 1, Meda 1, and Meda 2 wells; 30, Tappers Inlet 1 well; 31, Grant Range 1 well; 32, Roebuck Bay 

1 well; 33, Matches Springs 1 well; 34, Pegasus 1 well; 35, Gingerah Hill 1 well; 36, Kempfield 1 well; 37, Frankenstein 1 well; 38, Kidson 1 well; 39, Gneudna 

Formation, Carnarvon Basin; 40, Munyarai 1 well, Officer Basin; 42, Grampians; 43, Point Hibbs; 44, Reefton, South Island, NZ; 46, Lorentz River, Irian Jaya; 47, 

Warsampson River, Kepala Burung, Irian Jaya; 48, Buldah (Vic.).

(Fig. 5H) and? Yealepis or Nostovicina guangxiensis (Fig. 5I) 

dominate the acanthodian fauna, with rare examples of poracanthodid 

fin spines (Fig. 5C), tooth whorls (Fig. 5D), and 

possible climatiid tooth whorls (Fig. 5E). The diversity of placoderms 

increased in the late Lochkovian, although the rich 

assemblages comprise only small individuals represented by 

scales, fragmentary dermal bone, and rare whole plates and 

eye ‘capsules’ (Burrow, 2006; Burrow et al., 2005). Placoderms 

include acanthothoracids comparable with the circumArctic 

Romundina (Fig. 5N), Palaeacanthaspis, Kosoraspis (Fig. 5O), 

and Radotina from the Czech Republic, and Hagiangella from 

Vietnam (Racheboeuf et al., 2005), as well as the endemic 

Connemarraspis and the basal buchanosteid Narrominaspis. 

Another noteworthy taxon found at this level is Lophosteus 

incrementus (Fig. 5T), represented by scales, spinal elements, 

tooth plates, and dermal plate fragments. Botella et al., 2007, 

and earlier workers, regard Lophosteus as a putative stem osteichthyan 

genus. Other scales and dermal bone fragments might 

represent stem coelacanths (Burrow, 2007; Fig. 5U), as they are 

ornamented similarly to scales on younger articulated coelacanths, 

but no definitive elements have yet been found. 

Burrow (2002, table 6) described the progression of acanthodian 

taxa through the Early Devonian of southeastern Australia. 

Other occurrences of vertebrate micro-remains include the?late 

Lochkovian/early Pragian Martins Well Limestone Member, 

Shield Creek Formation of the Broken River region, north 

Queensland (Turner et al., 2000, table 1). The acanthodian 

taxa listed show a closer resemblance to assemblages of similar 

age from the Taimyr region of Russia (Valiukevičius, 1994) 

than to those of southeastern Australia, although the?endemic 

Garralepis simplex Burrow, 2002 is found in both Australian 

regions. Carbonates with conodont and microvertebrate faunas 

of mid-Pragian age are rare in Australia. The only undoubted 

kindlei CZ vertebrates are from the Coopers Creek Limestone 

in the Tyers-Boola region of Victoria (Basden, 1999a), with a 

diverse assemblage incorporating acanthodian taxa also known 

from older (T. cervicostulata, G. simplex) and younger deposits 

(Radioporacanthodes liujingensis, Gomphonchus? bogongensis, 

Nostolepoides platymarginata). Mawson et al. (1988) 

suggested the lack of conodont-producing carbonates from 

kindlei-pirenae CZ strata could be related to a regressive interval 

in eastern Australia. A rich assemblage of vertebrate microremains 

in the mid-late Pragian Fairy Formation of Victoria 

includes a jaw fragment of the oldest known coelacanth Eoactinista 

foreyi Johanson et al., 2006, and a new onychodont, 

Bukkanodus jesseni Johanson et al., 2007 (Johanson et al., 

2006, 2007) (also one of the oldest known, and its presence 

first noted by Turner, 1991). Vertebrate macro-remains 

were first recorded from the overlying, early Emsian Buchan 

Group in the 1870s; micro-remains were not fully described 

until the work by Basden (2003). The assemblages compare 

closely with those from the Taemas/Wee Jasper region in New 

South Wales and the late Pragian-early Emsian Gleninga Formation 

of central western New South Wales (Basden, 1999b; 

Basden et al., 2006; Burrow, 1997, 2002). Common elements 

include scales of acanthodians N. platymarginata, N. guangxiensis, 

G.? bogongensis, and Cheiracanthoides spp., arthrodire 


placoderms Goodradigbeeon and Buchanosteus, petalichthyid 

placoderm Lunaspis sp. (Fig. 5Q), acanthothoracid placoderm 

Murrindalaspis sp. (Fig. 5R),?chondrichthyan Ohiolepis sp., 

actinopterygian Ligulalepis toombsi, and onychodontid scales 

and teeth. The early-mid Emsian marks the incoming of several 

new acanthodian taxa, Gomphonchus? bischoffi (Fig. 5L) 

and Gomphonchus? fromensis (Fig. 5M) at Murrindal, Victoria, 

and several southeastern-central New South Wales regions 

(Burrow, 2002) including recent recordings from the Bongalaby 

Formation near Braidwood (CJB, pers. obs., 2008). 

Recent investigations of microvertebrate assemblages elsewhere 

have shown a more widespread distribution of some of the 

late Pragian-early Emsian acanthodians and placoderms found in 

marine limestones of eastern Australia and New Zealand, where 

Turinia sp. occurs with placoderm and acanthodian remains 

in the Reefton Limestone (Macadie, 2002). Acanthodian G.? 

fromensis (Fig. 5M) and an acanthothoracid with scales identical 

to those of Jerulalepis picketti (Fig. 5P), only known in Australia 

from the Gleninga Formation (Burrow, 1996), also occurs in the 

early Emsian Jawf Formation of Saudi Arabia (Burrow et al., 

2006), and R. liujingensis (Wang, 1992) is identified in the early 

Emsian of Uzbekistan (Burrow et al., 2008a, 2010) as well as in 

the Pragian-early Emsian of China and Australia. Canadalepis 

basdenae Burrow, 2002, now recognized in the early Emsian 

of the Zinzalban section, Uzbekistan and also possibly the Jawf 

Formation, occurs at higher levels (serotinus CZ) in eastern Australia. 

The placoderm scale taxon Xiejiawangaspis Burrow et al., 

2000, now known from the Sichuan Province, China and Uzbekistan, 

closely resembles scales of Murrindalaspis (Fig. 5R) from 

coeval southeastern Australian localities. Several scale-based 

acanthodian species of late Pragian-Emsian age which Burrow 

(1997, 2002) assigned to G.? bogongensis, G? fromensis, and G? 

bischoffi are probably from the ischnacanthiform fish to which 

the dentigerous jaw bone-based taxa Rockycampacanthus and 

Taemasacanthus spp. (e.g., Long, 1986; Lindley, 2000, 2002a,b) 

belong. 

In the Broken River area of north Queensland, the youngest 

strata of the Shield Creek Formation are mid-Pragian (kindlei 

CZ), with a depositional hiatus to the oldest late Emsian (inversus/laticostatus 

CZ) strata of the Broken River Group. The 

Group ranges up to the earliest Frasnian, with rich vertebrate 

micro- and macroremain assemblages found at all levels (Turner 

et al., 2000), some still awaiting full description. 

By assuming a Silurian age for the Silverband Formation, the 

oldest Devonian thelodont scales, assigned by Turner (in Pickett 

et al., 1985) toTurinia cf. polita, occur in the?late Lochkovian 

Tumblong Oolite. Turinia sp. is found in the upper levels of 

the Connemarra Formation, which are probably of early Pragian 

age and also in the Broken River Martins Well Limestone, 

northern Queensland and certain beds of the Cravens Peak Beds, 

northwestern Queensland (e.g., Turner et al., 2000). 

Revision of the dating for strata in the Trundle Group of 

central New South Wales (e.g., Sherwin, 1996), supported by 

the composition of the gnathostome micro-remain assemblages, 

indicates that all occurrences of Turinia australiensis Gross, 

1971 sensu stricto, both marine and non-marine, are of late 

Pragian-early Emsian age (contra Turner, 1997a, fig. 2). The


Fig. 5. Early-Middle Devonian vertebrate micro-remains of biogeographic significance from Australia. (A and B) Thelodont agnathan scales. (A) ANU V3457, 

Turinia cf. hutkensis, lateral view; (B) ANU V3455, T. gavinyoungi, anterocrown view. (C–M) Acanthodian elements. (C) MMMC04410, poracanthodid fin spine; (D) 

MMMC04396, tooth whorl with central row of large sharp teeth and fused lateral rows with smaller teeth, occlusal view; (E) MMMC04397, tooth whorl of blade-like

distribution of T. australiensis extends from the marine limestones 

of south-eastern Australia, through the predominantly 

non-marine Darling and Amadeus basins westward. Although 

one of us (GCY) regards these basins as non-marine freshwater, 

the turiniid records indicate periodic shallow water marine 

incursions, possibly indicative of hyper-, or hyposaline, lagoonal 

environments. 

3.2.2. Non-marine-marginal marine assemblages 

Distribution of the thelodont T. australiensis and closely 

related species extends westward from the Mulga Downs Group, 

Darling Basin, western New South Wales, to the N’Dahla Member 

of the Pertnjara Group, Amadeus Basin, central Australia 

(Young et al., 1987; Fig. 5A), on to the type locality of Wilson 

Cliffs in the Canning Basin, and other boreholes in Western 

Australia (Gross, 1971; Turner, 1995 [redescription and SEMs 

of the type material], 1997a). Identification of these occurrences 

as the same, or closely related, species is supported by the cooccurrence 

of two acanthodian species at all these localities; 

unfortunately the scales of these acanthodians, unlike those of 

the thelodont, are very rare, and differ from all described species 

particularly in their histological structures (cf. Gross, 1971, figs. 

2 and 3; Burrow, 2002, fig. 13A–C, 14J and K). Acanthodian 

Striacanthus fin spines from the Mulga Downs Group are diplacanthiform 

(Burrow, 2002), and presumably are from the same 

fish as the scales with Diplacanthus-type structure (Gross, 1971, 

fig. 2A). The other scale type (Fig. 5J) appears to be from a new 

genus. 

As detailed by Young et al. (2010), opinions differ about 

the age of these assemblages, from Pragian to early Eifelian. A 

late Emsian-Eifelian age for the Cravens Peak limestone assemblage 

is supported by the occurrence of the thelodont Turinia 

gavinyoungi Turner, 1995 at this locality and also in well-dated 

marine limestones from this time interval in the Broken River 

Group. This taxon also resembles those of a similar age across 

Gondwana including China and Iran (Turner, 1997a). The small 

Cravens Peak limestone outcrop has yielded a rich and very 

well-preserved assemblage of vertebrate micro-remains as well 

as rare jaw fragments and plates. New work has been published 

on the shark Mcmurdodus whitei Turner et Young, 1987 (Turner 

and Young, 1987; Burrow et al., 2008b; Fig. 5S), acanthodians 

Terenolepis toombaensis and Machaeracanthus pectinatus 

(Burrow and Young, 2005), and osteolepid, holoptychiid, dip- 


noan, and onychodontid (Fig. 5V) osteichthyans (Young and 

Schultze, 2005). 

The? non-marine Hatchery Creek Formation at Wee Jasper, 

conformable on the Emsian limestones, of assumed Eifelian 

age (Young and Gorter, 1981), has a completely different fish 

assemblage, which includes the thelodont Turinia cf. hutkensis 

(see Turner, 1997a), plus various endemic taxa. The association 

of bothriolepids (Monarolepis) with turiniid thelodonts, 

also documented in other Gondwanan sequences (e.g., Antarctic 

Aztec fauna, Young, 1988; Gneudna fauna of Western Australia, 

Long and Trinajstic, 2000; Chariseh fauna, Hairapetian 

et al., 2006), has never been recorded from northern hemisphere 

blocks. The persistence of turiniid thelodonts into the 

late Frasnian (Turner, 1997a; Trinajstic, 2000) is characteristic 

of Gondwanan sequences only. 

3.3. Middle–Late Devonian (Givetian-Famennian) 


Marine fish faunas in the Givetian-Frasnian interval include 

both endemics and widespread forms. Turner et al. (2000, table 

2) and Young and Turner (2000) summarized the distribution 

of vertebrate micro-remains from the Broken River region in 

Queensland. Rare but significant are the key Palaeotethyan cosmopolitan 

chondrichthyan taxa Phoebodus, Protacrodus, and 

Stethacanthus (e.g., Jones et al., 2000b; Turner et al., 2000; 

Young and Turner, 2000). 

Basden et al. (2006) summarized new work done by 

Trinajstic (e.g., 2000, 2001a,b) on the species-rich microvertebrate 

assemblages from the Frasnian Gneudna Formation, 

Western Australia, which include scales and other elements 

from thelodonts Australolepis seddoni and Turinia hutkensis, 

placoderm Holonema westolli, palaeoniscoids Moythomasia 

durgaringa and Mimia toombsi, acanthodians, chondrichthyans, 

and sarcopterygians. Of note too is increasing evidence of links 

in thelodont, placoderm, chondrichthyan, and actinopterygian 

taxa between the Frasnian to Famennian of Western Australia 

and Iran (e.g., Turner et al., 2002; Hairapetian and Turner, 2003; 

Turner and Hairapetian, 2005; Hairapetian et al., 2006). 

3.3.2. Non-marine-marginal assemblages 

An outcrop of calcareous grey siltstone in the karawaka biozone 

(?late Givetian) of the Aztec Siltstone at Mt. Crean in 

teeth with squat sharp central cusps, occlusolateral view; (F) AMF97973, Zemlyacanthus menneri scale, crown view; (G) MMMC04398, Radioporacanthodes 

sp., anterocrown view; (H) MMMC04399, Nostovicina guangxiensis, crown view; (I) MMMC04400, Trundlelepis cervicostulata, crown view; (J) ANU V3458, 

Acanthodii n. gen. n. sp., anterior view; (K) MMMC04395, Cheiracanthoides wangi, crown view; (L) MMMC04394, Gomphonchus? bischoffi, anterocrown 

view; (M) ANU V3459, Gomphonchus? fromei, crown view. (N) Romundina sp. Reconstruction showing arrangement of premedian (MMMC04401), suborbital 

(MMMC04402), preorbital (MMMC04403), fused postorbital+marginal+anterior paranuchal (MMMC04404) plates, with sclerotic capsule (MMMC04405). (O) 

MMMC04406, Kosoraspis? sp. posterior dorsolateral plate. (P) MMMC02299, Jerulalepis picketti scale, crown view. (Q) MMMC04409, Lunaspis sp. spinal plate, 

distal end. (R) ANU V1638.12, Murrindalaspis sp. scale, dorsal view. (S) QMF52817, Mcmurdodus whitei tooth, labiobasal view. (T) MMMC04407, Lophosteus 

incrementus dermal bone fragment, lateral view. (U) MMMC04408, possible coelacanth scale, closeup of posterior half. (V) ANU V3456, Luckeus abbuda, occlusal 

view of parasymphysial tooth whorl, main cusps broken off. (A and J) From sample ND5, N’Dhala Mb. (?late Emsian), Pertnjara Gp, locality 17, Fig. 4; (B, S, and V) 

from sample GY7, Cravens Peak Beds, limestone outcrop (?late Emsian-early Eifelian) south end of locality 15, Fig. 4; (C–D, F–I, N, O, T and U) from GSNSW site 

C669, Connemarra Fm. (?late Lochkovian), locality 10, Fig. 4; (E) from Camelford Lst., earliest Lochkovian, locality 5, Fig. 4; (K and L) from GSNSW site C2047, 

Bongalaby Fm. (early Emsian), locality 5, Fig. 4; (M) from?Bloomfield Lst. (early-middle Emsian), Murrumbidgee Gp., locality 5, Fig. 4; (P) from GSNSW site 

C231, Jerula Mb. (Pragian/Emsian boundary), Gleninga Fm., locality 10, Fig. 4; (Q) from GSNSW site C595, Troffs Fm., locality 10, Fig. 4; (R) from Murrumbidgee 

Gp., Taemas, locality 5, Fig. 4. Scale bar = 0.01 mm in (B), 0.1 mm in (A, C, E–M, O–Q and T–V); 1 mm in (D, N, R and S).


southern Victoria Land, Antarctica yielded a rich microvertebrate 

assemblage (Burrow et al., 2009) comprising acanthodians 

Pechoralepis juozasi, Nostolepis cf. gaujensis, Milesacanthus 

antarctica, and an undetermined acanthodiform, as well as 

chondrichthyan scales and teeth (Antarctilamna and Aztecodus), 

thelodont Turinia antarctica scales, and palaeoniscoid scales and 

other elements. The acanthodian assemblage resembles those 

from early Frasnian carbonates of central Iran (Hairapetian et al., 

2006). Ginter et al. (2006) confirmed the synonymy of Antarctilamna 

and Wellerodus first mooted by Turner (1997b), thus 

extending the known geographical range of this genus and also 

Portalodus, both originally described from the Aztec Siltstone 

of Antarctica, to Laurussia (New York State). ‘Wellerodus’ has 

now also been recorded from the late Frasnian of the Middle 

Urals (Ivanov, 2008). 

A few chondrichthyan scales (Figs. 6A–C, 7A–D) and one 

of Acanthodes cf. australis (Turner et al., 1994; Fig. 7E–G) 

were retrieved from the Cann River Beds (Unit 3) of Buldah, 

in the Buldah-Club Terrace Belt, East Gippsland Province, Victoria 

(Marsden in Douglas and Ferguson, 1976, p. 119). The 

shark scales resemble those of a hybodontid/sphenacanthid and 

these plus the acanthodian support a late Famennian age; the 

palaeoenvironment might be marginal marine. The oldest known 

species of Acanthodes based on articulated specimens is A. lopatini, 

from the Tournaisian of Russia (Beznosov, 2009). While 

Acanthodes-type scales are recorded from many Middle–Late 

Devonian assemblages, their assignment to Acanthodes s.s. must 

remain doubtful. 

3.4. D/C boundary assemblages 

As discussed elsewhere, locating the Devonian/Carboniferous 

boundary in Australia is a frustrating 

business especially in the various basins dominated by nonmarine 

sequences. A trench in the Oscar Hill area, Fitzroy 

Trough, Canning Basin of Western Australia through the 

boundary beds yielded only conodonts and no vertebrates from 

the Famennian (J. Talent, pers. comm.), but the Tournaisian 

part produced a good marine vertebrate fauna (Edwards, 1997) 

comprising chondrichthyan teeth and actinopterygian teeth and 

scales (see Section 4.2.1). 

The non-marine-marginal redbeds of the Devil’s Plain Formation 

in the Broken River district near Mansfield, Victoria, 

dated as Tournaisian for much of the 20th century, were thought 

by the first worker, George Sweet in the 1890s, to be Devonian 

based on both their colour and geological setting; the 

general fauna and flora do not discount this possibility. Warren 

et al. (2000) redescribed Gyracanthides murrayi from Mansfield, 

recognizing the distinctive endoskeletal scapulocoracoid 

and procoracoid structures and figuring the distinctive spiny 

odontode-bearing scales in the specimens. Garvey and Turner 

(2006) supported a possible latest Famennian date for the Mansfield 

fauna of Victoria based on lithology and presence of 

faunal taxa (Gyracanthides, Ageleodus, xenacanthiform) known 

in the late Famennian of Pennsylvania and elsewhere (Turner 

et al., 2005), supporting the hypothesis of post-collision dispersal 

between East Gondwana to Euramerica (Laurentia) via 

northwest Gondwana (where Gyracanthides is known only from 

Iran). The Home Station Sandstone Member of the Snowy Plains 

Formation, Mansfield Group (Vandenberg et al., 2006) at Mansfield 

has yielded two plesiomorphic ‘enigmatic’ rhizodonts, the 

large Barameda decipiens and the small B. mitchelli (Holland 

et al., 2007), which might support Garvey and Turner’s (2006) 

assessment of the age of the Mansfield sequence as spanning 

the D/C boundary rather than being only Early Carboniferous. 

In addition, Mansfield has provided no tetrapods as yet, despite 

much searching. The only negative to this hypothesis is the 

absence of placoderm remains. 

3.5. Palaeobiogeographic significance (Devonian 

microvertebrates) 

In general for the diverse Devonian record, many taxa are 

now considered to be endemic. There are, however, strong 

and increasing links throughout the period to neighbours and 

across North Gondwana (Iran, Middle East, South America, 

Spain); eventually as landmasses approached collision, faunal 

links (e.g., chondrichthyans, gyracanthids, tetrapods) arose associated 

with the Appalachian orogen of Laurentia. Taxa were 

living in primarily equatorial to warm temperate settings, as in 

earlier periods, with indications of speciation related to transgression/regression 

across areas of Australia. 

4. Carboniferous vertebrates (see Fig. 1 for localities) 


Australasian marine invertebrate faunas from the early Carboniferous 

indicate warm shallow seas, with the craton at a low 

latitude between 0 ◦ and 40 ◦ south. The rapid loss of faunal and 

floral diversity around the Viséan-Namurian boundary marks the 

beginning of the southern hemisphere ice age accompanied by 

the movement of eastern Gondwana to high latitudes (Jones and 

Metcalfe in Jones et al., 2000a; Fielding et al., 2008). 

4.2. Mississippian vertebrates 


Ichthyoliths were recovered from a 450-m trench excavated 

by a Macquarie University crew across the Devonian- 

Carboniferous boundary and extending some way above it 

into the Tournaisian near Linesman Creek, and some isolated 

outcrops in the Oscar Hill area, north Canning Basin, 

Western Australia. Remains from the early Carboniferous Laurel 

and Gumhole formations comprise fish teeth, scales, and 

bones from several orders, mirroring earlier studies (e.g., 

Turner, 1991): most abundant are those of the Actinopterygii 

(palaeoniscoid dermal bones, teeth and scales) and Acanthodii 

(Acanthodes scales and spines), and one stellate tubercle that 

could be from a shark or acanthodian. Sarcopterygian teeth 

and scales are also present. By far the greatest taxonomic 

diversity lies within the Chondrichthyes, with representatives 

of families Protacrodontidae, Ctenacanthidae, Phoebodontidae, 

Stethacanthidae, Orodontidae, Hybodontidae, Holocephali, and


Fig. 6. Late Devonian-Carboniferous vertebrates from Australia. (A–C) Shark scales from the?Famennian Cann River Beds (Unit 3), Victoria, locality 48, Fig. 4.(A 

and B) NMV P229481, lateral views; (C) NMV P229482, basal view. (D–J) Microvertebrates from the Viséan Utting Calcarenite (Chambers, 2002), locality C6 (QML 

1095). (D) Hybodontiform shark tooth, lingual view (tooth lost in SEM); (E) QMF48858, protacrodont shark tooth,?labial view; (F) Thrinacodus? sp. tooth, lingual 

view (tooth lost in SEM); (G) QMF48920, Mesodmodus? sp. tooth, labial view; (H) QMF49054, bradyodont cf. Diclitodus sp. tooth,?lingual view; (I) QMF49059, 

possible shark head or clasper denticle; (J) QMF49005, palaeoniscoid palatal teeth. (K–M) Shark teeth from the Kyndalyn Mb., Merlewood Fm., locality C2. (K) 

ML-1, symmoriid, occlusolabial view; (L) ML-2, Lisssodus? sp., lingual view; (M) ML-3, caseodontid, occlusal view. (N) Largest of three blocks of the nodule 

ANU V1772 from the Utting Calcarenite, locality C6, with prismatic calcified cartilage jaw segment and large teeth of a stethacanthid shark, probably S. thomasi,a 

unique find so far in Australia. (O–R) Chondrichthyan and other elements from the late Viséan Baywulla Fm., locality C4. (O) QMF54772, Bransonella sp. tooth, 

labial view; (P) QMF54772,?neoselachian scale, crown view; (Q) QMF54773, hybodontiform scale, anterior view; (R) QMF54774,?sarcopterygian harpoon-shaped 

tooth cf. Hamodus. (S) UQY F73008, xenacanthiform tooth from the Early Carboniferous of the Broken River Gp., BRJ 50 of Turner et al. (2000), locality C8. (T 

and U) Tetrapod Ossinodus pueri scutes/scales, in matrix with specimens outlined, from the Ducabrook Fm., locality C5. (T) QMF37517, internal surface showing 

overlap corner, whitened with magnesium oxide; (U) QMF37515, external surface showing lightly ornamented bone. Scale bar = 0.1 mm in (A–D, L–M), 0.2 mmin 

(E–G, K and Q), 0.5 mm in (H and R), 1 mm in (I, J, O, P and S); 1 cm in (N, T and U).


Fig. 7. Drawings (by ST) of some Late Devonian to Carboniferous specimens. (A–G) Microvertebrates from the?Famennian Cann River Beds, locality 48, Fig. 4. 

(A–D) Hybodontid/sphenacanthid shark scale NMV P229483; (E–G) acanthodiform acanthodian scale NMV P229484 in crown, basal, and lateral views. (H) ANU 

V1772, Utting Calcarenite nodule, broken into three blocks, with stethacanthid jaw cartilage and displaced teeth. cc, prismatic calcified cartilage. Scale bar = 0.1 mm 

in (A–G); 2 cm in (H). 

Paraselachii (Edwards, 1997). Protacrodonts include Protacrodus 

wellsi, Deihim, and cf. Orodus decussatus—teeth assigned 

to this genus are typical of Devonian-Carboniferous boundary 

beds in Australia, China, USA, Germany, and elsewhere 

(e.g., Ginter, 1999, Thuringia); phoebodontids include Jalodus 

australiensis, Thrinacodus ferox, and Thrinacodus bicuspidatus 

Ginter et Sun, 2007 (Ginter and Sun, 2007), well known 

as cosmopolitan Tethyan sharks from Australia, China, Ireland, 

UK, and USA (e.g., Blieck and Turner, 2000); several stethacanthids 

include Stethacanthus sp. cf. Stethacanthus thomasi 

and cf. Denaea sp.; hybodontids include Lissodus; other chondrichthyans 

are cf. Venustodus and a helodont. Studies of the 

assemblages in the Mississippian limestones of the Canning 

Basin await further description (Turner, 1993b; Trinajstic and 

George, 2007). Several taxa can be correlated with those from 

the Devonian-Carboniferous boundary deposits elsewhere in 

Australia, SW Asia, North America, and Europe. 

Two predominantly holocephalan faunas have been studied 

thus far: a series of records from the Tournaisian to 

Viséan in Queensland (Turner, 1990); and one from the Viséan 

Utting Calcarenite, Weaber Group, Bonaparte Basin, northern 

Western Australia (Turner in Jones et al., 2000a; Chambers, 

2002; Trinajstic, Turner and Chambers, in prep.); both contain 

many higher taxa in common with those worldwide but 

evidence of endemic species is coming to light (Fig. 6D–J). 

The Utting Calcarenite is a highly fossiliferous unit that contains 

an excellently preserved collection of vertebrate macroand 

microfossils, principally symmoriid and durophagous chondrichthyan 

teeth, palaeoniscoid scales and a sarcopterygian, plus 

a diverse assemblage of marine invertebrates. Taxa comprise

at least 18 different chondrichthyans: Helodus spp., Psephodus; 

cochliodonts; cf. Poecilodus; cf. Venustodus/Mesodmodus; 

Psammodus; protacrodonts; Orodus; petalodonts; Thrinacodus; 

ctenacanths; Symmorium; stethacanths; denaeid; and Lissodus. 

This formation had yielded a large ctenacanth spine and notably 

one half of a nodule (ca. 10 cm × 10 cm × 5 cm), first mentioned 

by Joyce Gilbert-Tomlinson (in Veevers and Roberts, 1968). 

This last specimen (featured in Long, 2006) contains the partial 

jaw of a large stethacanthid shark, with ca. 20 teeth in situ in 

jaw cartilages with associated external dermal scales and mucous 

membrane scales from inside the mouth (Turner, 1991; Turner in 

Jones et al., 2000a; Turner et al., 1994), thus allowing the identification 

of scales with associated teeth (Figs. 6N and 7H). Unfortunately 

the counterpart is unknown and no other similar specimens 

have been found in the area in 50 years, but clearly some 

very large sharks were inhabiting the reefal areas in Viséan times. 

In NSW, the Bingleburra, Namoi, Merlewood (Kyndalyn 

Member) and Dangarfield formations have yielded Thrinacodus, 

ctenacanthoid, stethacanthid (Fig. 6K–M), hybodontids, and 

eugeneodontids such as caseodontid teeth and even neoselachian 

placoid scales correlatable with faunas in China, Europe, and 

North America (Turner et al., 1994; Turner, Jones and Leu, in 

prep.). 

4.2.2. Non-marine assemblages 

Much work has been done on bone bed assemblages from 

the Drummond Basin, especially on the M99 bonebed from 

the Narrien Range first described by Turner (1993c); over the 

last decade, students at Monash University assisted in picking 

residues from this bonebed and have brought to light 

further chondrichthyans (xenacanth, hybodontiform, polyrhizodont) 

and 5–6 taxa of palaeoniscoids and a megalichthyid 

sarcopterygian, mainly represented by scales, teeth, and dermal 

bones; this wider study has yet to be published. Fox et 

al. (1995) described articulated specimens of the large osteolepiform 

megalichthyid Cladarosymblema narrienense and 

Long (1986) figured acanthodiform acanthodian jaws from 

the Raymond Formation. Xenacanths and other sharks from 

these Tournaisian to early Viséan bonebeds are being described 

(Turner et al., 2008; Turner and Burrow, in prep.). 

Carboniferous work during the project time span has centred 

on a fauna of fish and tetrapods from the Early Carboniferous 

(mid-Viséan) Ducabrook Formation in the Drummond Basin, 

Queensland (Warren and Turner, 2004). This, the only Gondwanan 

Carboniferous fauna containing tetrapods and dated as 

Holkerian-Asbian, falls near the end of Romer’s Gap, a 25million 

year period following the end of the Devonian in which 

tetrapod fossils are rare. The Ducabrook fauna is the only tetrapod 

fauna in Australia between the Late Devonian and near 

the end of the Permian, a span of some 100 million years. A 

single stem tetrapod taxon, Ossinodus pueri Warren et Turner, 

2004, has been described from this fauna; the find in 2004 of 

most of a complete portion of the tetrapod skull, a dorsal skull 

roof extending from the external nostril to back of the quadrate 

allowed further refinement of Ossinodus (Warren, 2007); unlike 

many younger tetrapods, Ossinodus had bony scutes/scales on 

the body (Fig. 6T and U). 


Fish from the Ducabrook Formation comprise large sarcopterygians: 

a rhizodont, Strepsodus sp. (Johanson et al., 2000; 

Parker et al., 2005), and a lungfish, Ctenodus sp. (Turner et al., 

1999). The dominant taxon in the assemblage is the gyracanthid 

acanthodian Gyracanthides hawkinsi (Turner et al., 2005), with 

close similarities to contemporaneous species across northern 

Gondwana in Ghana (Mensah and Smit, 1972), and in the thencollided 

Laurentian parts of Scotland, maritime Canada, and 

the eastern Appalachians. Several palaeoniscoid actinopterygians 

are known from ornamented bones and scales but are 

not yet described. Chondrichthyans, present as microvertebrates 

(Turner et al., 1996), include xenacanthiforms (teeth, spines, 

and possible jaw), hybodontiforms, and Ageleodus (Garvey and 

Turner, 2006; Turner, in prep.). The new xenacanth adds a further 

dimension to the early evolution of xenacanthoid sharks in Gondwana, 

which appear first in the latest Devonian (Mansfield), 

Tournaisian (Narrien Range, Drummond Basin) and then Viséan 

(Drummond Basin) (Turner et al., 2008); a solitary xenacanth 

tooth is still all that is known from the Early Carboniferous of 

the Broken River region (Fig. 6S). 

Another localized occurrence of a similar age to the 

Ducabrook assemblage is found in the Bulliwallah Formation, 

northern Drummond Basin. The fossil fish are preserved, usually 

as isolated elements, in small siltstone nodules (Burrow and 

Turner, 2002); taxa include actinopterygians, sarcopterygians, 

Gyracanthides, and the acanthodiform Acanthodopsis russelli 

Burrow, 2004 (Burrow, 2004). The study of Australian gyracanthid 

acanthodian remains prompted a review of all known taxa 

across Gondwana that has differentiated Early Devonian from 

younger taxa (Burrow et al., 2008c). 

No more tetrapods are found until the Late Permian presumably 

because of the onset of the southern hemisphere Ice Age 

in the mid-Carboniferous. Fielding et al. (2008), however, have 

shown the oscillating effect of glacial conditions in eastern Australia/Gondwana/Pangaea 

with the warmer interglacial periods 

allowing for some development of reefal or similar environments 

at least in parts (Queensland). 

4.3. Younger Carboniferous 

Remaining vertebrate microfaunas, all from Queensland 

where environments were still vertebrate-friendly (Turner et al., 

1994; Turner in Jones et al., 2000a), are not yet fully described. 

One small sample from the Baywulla Formation includes a tooth 

of Bransonella (Fig. 6O), well known now in Tethyan localities 

in Eurasia (e.g., Timan, and possibly Iran and China: Ginter et 

al., 2002), along with possible hybodont and neoselachian scales 

and at least one unusual sarcopterygian tooth (Fig. 6P–R). 

The last known assemblage from the Barambah Limestone 

at Murgon (Turner, 1993a) represents the youngest in the Carboniferous, 

of Pennsylvanian age and closest to denaeid shark 

material described from the USA by Williams (1985). 

4.4. Palaeobiogeographic significance (Carboniferous) 

In eastern Australia, the Late Palaeozoic Ice Age was represented 

by discrete glaciations, separated in time by intervals


of nonglacial conditions (Fielding et al., 2008). The fossil vertebrate 

record in the Mississippian to Pennsylvanian in NSW 

and particularly Queensland provides indicators of probable 

interglacial conditions, with its series of chondrichthyanpalaeoniscoid 

assemblages from limestones and bonebeds, 

especially the non-marine xenacanth-hybodontiform-tetrapod 

faunas. The only Carboniferous tetrapod fauna in the southern 

hemisphere (mid-Viséan Ducabrook Formation) and the 

youngest Carboniferous assemblage that occurs in southern 

Queensland in the mid-Pennsylvanian (Barambah Limestone) 

support the findings by Fielding et al. (2008) that glaciation was 

not so severe in eastern Australia at that time. 

5. Discussion and Conclusions 

We are slowly refining the timing of the reappearance of vertebrates 

after the Late Ordovician-earliest Silurian Ice Age. The 

discovery of an important Ordovician fauna in Oman (Sansom 

et al., 2009) prompts the need for further studies across Gondwana. 

Interestingly, Neef (2007) noted similar environmental 

conditions in Oman and Eastern Australia producing alluvial-fan 

deposits. 

A closer look for more Gondwanan Silurian thelodonts, acanthodians, 

placoderms, and early sharks should be a priority for 

the future, bearing in mind the relationship of these with Iran and 

parts of China. We need to learn more about possible endemic 

and/or difficult to explain taxa, such as those from Irian Jaya 

and the unique Mcmurdodus. Links between Australian regions 

and other terranes, and within Australian regions, vary markedly 

over time. As noted earlier, the greatest such variations in affinities 

occur during the Silurian, despite the poor vertebrate record 

for this period. 

In contrast to the (northern hemisphere) Laurentia terranes, 

of particular interest are the diversity and radiation of turiniid 

thelodonts in Gondwana, including taxa which outlasted the 

other major agnathan groups into the latest Frasnian and Famennian, 

with Western Australia and Iran as refugia for turiniids 

(Trinajstic, 2001a; Turner et al., 2002, 2004; Hairapetian and 

Turner, 2003; Turner and Hairapetian, 2005). Interestingly, also, 

whereas there is a strong affinity between East and West Gondwanan 

thelodonts in the Early to Middle Devonian (e.g., T. 

gondwana, T. antarctica, T. hutkensis, T. gavinyoungi and T. spp. 

in South China, West Yunnan, central Australia and Thailand), 

turiniids then disappear from the presumed cooler to cold southern 

Malvinokaffric faunas with their endemic chondrichthyans 

and acanthodians (cf. Turner, 1997a, fig. 9). 

By the end of the Devonian, many chondrichthyan taxa show 

Palaeotethyan distributions, especially in the post-F/F radiation 

of sharks: Thrinacodus, Jalodus, phoebodonts, protacrodonts, 

symmoriids, stethacanths, ctenacanths with hybodontiforms and 

even neoselachian taxa appearing or on the increase. The major 

radiation with incoming of durophagous ‘bradyodont’ forms 

takes off in the Early Carboniferous after placoderms disappear, 

although such durophagous forms co-occurred with placoderms 

as early as the Middle Devonian (Givetian) in Euramerica 

(Darras et al., 2008). 

Of great import still is the interchange of taxa after 

the Gondwana-Laurentia collision, with a distinctive, occasionally 

tetrapod-bearing, assemblage (Ageleodus, hybodonts, 

xenacanths, ctenodont lungfish, rhizodonts, gyracanths, acanthodiforms) 

characterising Late Devonian into Mississippian 

non-marine settings and associated with rapidly expanding lepidodendroid 

forests. 

Bonebeds are spasmodic in the Silurian to Carboniferous 

record in Australia; most probably represent tempestites in 

deltaic and estuarine settings, especially in the Early Carboniferous 

of the extensive Mississippi-like Drummond Basin. 

Increasing aridity (even associated with glacial conditions) probably 

prompted droughts and flooding (equivalent to El Niño-La 

Niña events in modern history) causing widespread fish (and 

tetrapod) mortality as seen in key localities in eastern Australia 

(see also Young et al., 2010). Some vertebrate bone 

beds in the Australasian geological record represent mass-death 

events (e.g., Canowindra, Ducabrook) whereas others (e.g., Narrien 

M99) provide examples of lag (Kondenzat-Lagerstätten) 

deposits. More attention to the taphonomic information from 

these sites should bring forth more palaeoclimatic information. 

Talent (1993) summed up nicely the achievements and 

predicament of the Australian mid-Palaeozoic (S-D-C) vertebrate 

work and needs when we began the cooperative IGCP 

328 project. Some of the questions he raised and suggestions 

he made have been realised during the life of IGCP 491. Much 

still remains to be done both taxonomically and in terms of geochemical 

work and other related tasks. Lack of funding both 

for such research and in Australia has blighted possibilities for 

palaeontologists in the life of IGCP 491. The major database and 

ability to check palaeogeographic data and hypotheses (planned 

initially in IGCP 328) is finally coming to fruition. The main 

problem now is the lack of students to undertake the work as Australia 

and the region still produce surprises with new localities 

and new taxa. 

Acknowledgments 

The authors acknowledge provision of facilities in their 

respective institutions, and the IUGS-UNESCO International 

Geoscience Program and Australian IGCP Committee for 

various levels of support for the duration of IGCP Project 

491. Funding by the Australian Research Council under 

ARC Discovery Grants DP0558499 and DP0772138 is gratefully 

acknowledged (GCY). GCY acknowledges support as 

an awardee of the Alexander von Humboldt Foundation 

(2000–2003), based in Berlin at the Institute für Paläontologie, 

Museum für Naturkunde, where Prof. H.-P. Schultze is thanked 

for provision of facilities, and K. Dietze, M. Otto, J. Pálfry, R. 

Soler-Gijon, E. Siebert, and others provided help and support; 

R. Parkes is thanked for providing specimen images. Many colleagues 

including P. Ahlberg, A. Blieck, H. Blom, G. Clément, 

V. Dupret, D. Goujet, P. Janvier, Z. Johanson, C. Klootwijk, O. 

Lebedev, H. Lelièvre, W.-J. Zhao and M. Zhu have discussed 

Devonian biogeographic and biostratigraphic problems over 

many years. ST thanks the Australian Academy of Science European 

Exchange grant (2006) and colleagues that enabled work at

the Institut für Geowissenschaften, Eberhard-Karls Universität 

Tübingen in 2006, 2007, and 2008 (as a Guest Researcher) and 

especially W.-E. Reif, J. Nebelsick, V. Benz and C.-D. Jung. We 

thank M. Zhu and A. Blieck for their helpful reviews. 

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