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Reptilia: Sauropterygia) Records of the Westem Australian Museum Supplement No. 57: 201-205 (1999). Recent developments in Australasian sauropterygian palaeontology (Reptilia: Sauropterygia) Arthur R.I. Cruickshankl, R. Ewan Fordyce2 and John A. Long3 1 Earth Sciences Section, Leicester City Museum, New Walk, Leicester LE1 7EA, UK, and Geology Department, Leicester University, Leicester LII 7RH, UK; email: [email protected] 2 Department of Geology, Otago University, PO Box 56, Dunedin, New Zealand }Western Australian Museum, Francis Street, Perth, WA 6000, Australia Abstract -A brief review of recent research into Australasian sauropterygians is given. The earliest record is an Anisian pachypleurosaur from South Island, New Zealand. An Early Jurassic record from Queensland, hitherto included in the Plesiosauria is shown to be the last known pachypleurosaur. A diverse record of Pliosauroidea is known from the time of the break-up of East Gondwana, in marginal rift sediments of Western Australia and South Australia, dated to the earliest Cretaceous. The record of the marine incursion into the Eromanga Basin, Queensland, contains Albian-age pliosaurs and elasmosaurs. The New Zealand plesiosaurian record contains mostly members of the 'long-necked' Plesiosauroidea, all from the Haumurian stage of the Campanian-Maastrichtian. One specimen represents a unique record of a cryptoclidid in Australasia, more closely related to the European Callovian Cryptoclidus than to the other late cryptoclidids known from South America and the Antarctic Peninsula. INTRODUCTION NEW ZEALAND MATERIAL The sauropterygians were an entirely aquatic Sauropterygians have been known for over a group of reptiles which trace their ancestry to the century from the Haumurian Stage of the Late Permian tangasaurids of Madagascar, and Campanian-Maastrichtian in South Island, New whose intermediate evolutionary stages, the Zealand (Welles and Gregg 1971; Hiller et al. 1997). Pachypleurosauridae and the Nothosauriformes, More recently, a rich tetrapod fauna, including are best displayed in the Middle Triassic of the sauropterygians and other marine reptiles, has been European Alpine (Tethyan) region (Brown 1981; recovered from horizons of much the same age Carroll 1987). The best-known of the from the Mangahouanga Stream site in the North sauropterygians are the Plesiosauria, the earliest Island by Or Joan Wiffen and her associates (Wiffen of which is Pistosaurus, also occurring in the and Moisley 1986; Molnar and Wiffen 1994; Wiffen European Tethyan realm, and of Anisian (Middle 1996; Long 1998). Fordyce (1991) comprehensively Triassic) age. A functional progression from reviewed the New Zealand marine vertebrate laterally undulatory swimming, through a record, and Long (1998) included an illustrated 'rowing' phase, to underwater 'flight', akin to that summary of the known New Zealand Mesozoic practised by living sealions, is demonstrated in the marine reptiles. sauropterygians, with the plesiosaurians showing Another recent discovery in the New Zealand full adaptation to underwater flight. The Anisian, as part of the extension of the Tethys, Plesiosauria comprises the plesiomorphic comprises remnants of a pachypleurosaur Pliosauroidea (here we follow Brown 1981 in sauropterygian (Figure 2A). The evidence is in the recognizing only a single family, the Pliosauridae: form of a single very well preserved posterior but other classifications recognize the additional dorsal or anterior sacral vertebra, and other bone families Polycotylidae and Brachauchenidae); and fragments, showing strong affinities with European the more derived Plesiosauroidea, obtaining three forms, except that it is much larger than the families: the Plesiosauridae, the Elasmosauridae described species. and the Cryptoclididae. The last two are well The plesiosauroids (Figure 2B,C) are represented represented in the Australasian realm (Figure I, by forms in both the Australian Albian and the New overleaf). Zealand Maastrichtian. The New Zealand 202 A.R.I. Cruickshank, R.E. Fordyce, l.A. Long Casuarina Beach Mangahouanga Stream Tuarangisaurus, pliosaurid (LK) elasmosaur. pliosaur~ Darwin Richmond region 0 Woolungasaurus Kronosaurus (EK) Mount Morgan 0 Coober Pedy Kabarri Leptocleidus sp. pachypleurosaur (EJ) Leptocleidus cryptoclidid? (EK) Brisbane clemai (EK) o Haumuri t'-- Bluff (LK) Geraldton elasmosaur? (MJ) Lightning R~dge Cimoliasaurus ?(EK)J Waipara River (U<) Mauisaurus, pliosaurid Adelaide Sydney a)----- Shag Point MelbOU~ cryptoclidid? Dandaragan, Andamooka (LK) Gingin elasmosaur Otway Group , \ Strzelecki elasmosaur pliosaur (EK) plesiosaur (EK) Group (LK) M V plesiosaur 500 km (EK) 250- km Figure 1 Maps showing major occurrences of sauropterygian fossils in Australia and New Zealand. EJ = Early Jurassic; MJ = Middle Jurassic; EK = Early Cretaceous; LK =Late Cretaceous. elasmosaurs are historically important, having been large southern cryptoclidids in the Late Cretaceous reported from the earliest days of geological Palaeopacific, which, alongside the numerical exploration of that country (Long 1998). Two genera importance of the elasmosaurs, indicates an ocean of predominate in these faunas; Mauisaurus, the more than minimal productivity at that time. historically important taxon, and Tuarangisaurus, Pliosauroids are also present in the Late collected by Joan Wiffen and colleagues (Wiffen and Cretaceous of New Zealand, based on generically Moisley 1986). The former is not well defined, being indeterminate postcranial material from Waipara based on a quantity of dissociated limb and other River and Haumuri Bluff districts described by postcranial material, whereas the latter is known Welles and Gregg (1971), and from the from a single skull and some uncertainly-associated Mangahouanga Stream site (Wiffen and Moisley neck vertebrae. Unfortunately there is no overlap of 1986). this material. However, re-examination of fossil holdings in the Canterbury Museum and further collection of new specimens may solve this problem AUSTRALIAN MATERIAL (Hiller et al. 1997). A review of the Mount Morgan 'plesiosaur' Whereas elasmosaurids (Figure 2C) are the (Bartholomai 1966; Molnar 1991) has been apparently common plesiosaurian group found in undertaken by one of us (A.R.I.c.). This difficult New Zealand, a recently recognized cryptoclidid material consists of negative moulds in a from Shag Point, North Otago, presents a novel set ferruginous sandstone, and comprises remains of of adaptations (Cruickshank and Fordyce 1996, several individuals. The age is constrained as Early 1998). The specimen is from very near the top of the Jurassic (Playford and Cornelius 1967). A close Maastrichtian sequence, and at 6.5 m overall length examination of the material indicates that it has is among the largest of known cryptoclidids. The pachypleurosaur affinities, and is not plesiosaurian. skull is proportionally larger than normal for a The redescription will indicate that in the plesiosauroid and has relatively large orbits and Australasian Tethys there was an early temporal openings. Cladistic analysis shows it to be representative of the Sauropterygia and (if the closer to Cryptoclidus (Figure 2B) than to the other above identification is correct), the latest known two austral genera reported to date Aristonectes and pachypleurosaur. Turneria (Cabrera 1941; Chatterjee and Small 1989). It Although the plesiosaurs are taken to be a typical appears that there were at least two radiations of component of marine faunas, several Jurassic and Australasian sauropterygians 203 A ~I.._--- B c E Figure 2 Representative skeletons of Mesozoic sauropterygian groups discussed in the text. The scale bar for A = 10 cm; for B-E = 1 m. A, a pachypleurosaur Pachyplelirosalirlls, Family Pachypleurosauridae. B, a short-necked plesiosaur Cryptoclidlis, Family Cryptoclididae. C, a long-necked plesiosaur HydrotherosalirllS, Family Elasmosauridae. D, a long-necked pliosaur Dolichorhynchops, Family Pliosauridae. E, a short-necked pliosaur KrOnOSalirllS, Family Pliosauridae. A-C, E are after Carroll (1987), D is after Williston (1902). 204 A.R.I. Cmickshank, R.E. Fordyce, J.A. Long Early Cretaceous specimens have been recovered vertebra, plus smaller elongate vertebrae belonging from 'non-marine' horizons (Cruickshank 1997). In to elasmosaurs are also known from the early Late the Early Cretaceous (Barremian/Valanginian) of Cretaceous Molecap Greensand, at Dandaragan, Western Australia a new species, Leptocleidus clemai, Western Australia, and have recently been has recently been described from sediments described by Long and Cruickshank (1998). Other containing fossilized wood and a dinosaur vertebra South Australian elasmosaurs include the (Long and Cruickshank 1996; Cruickshank and 'Andamooka sea monster' (Rich and Rich 1985) and Long 1997). The genus Leptocleidus is known from a more unassigned opalized specimens, as well as close-inshore deposit in South Africa, and from a several occurrences reported and figured by Long non-marine 'Wealden' locality in England (1998) from isolated finds over many years. (Andrews 1911, 1922). Another closely related specimen is the famous 'Eric', an almost complete opalized skeleton from the Maree Formation, a CONCLUSIONS near-shore marine deposit at Coober Pedy in South The Australasian realm seems to have been an Australia, which has been the subject of a thesis important area for the evolution of leptocleidid recently submitted by Schroeder (1998). Other pliosaurs in the shallow seas which resulted from fragmentary
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