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Theropod Fauna from Southern Australia Indicates High Polar Diversity and Climate-Driven Dinosaur Provinciality

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Theropod Fauna from Southern Australia Indicates High Polar Diversity and Climate-Driven Dinosaur Provinciality Theropod Fauna from Southern Australia Indicates High Polar Diversity and Climate-Driven Dinosaur Provinciality Roger B. J. Benson1,2*, Thomas H. Rich3,4, Patricia Vickers-Rich3,4, Mike Hall3 1 Department of Earth Sciences, University of Cambridge, Cambridge, United Kingdom, 2 Department of Earth Sciences, University College London, London, United Kingdom, 3 School of Geosciences, Monash University, Melbourne, Australia, 4 Palaeontology Department, Museum Victoria, Melbourne, Australia Abstract The Early Cretaceous fauna of Victoria, Australia, provides unique data on the composition of high latitude southern hemisphere dinosaurs. We describe and review theropod dinosaur postcranial remains from the Aptian–Albian Otway and Strzelecki groups, based on at least 37 isolated bones, and more than 90 teeth from the Flat Rocks locality. Several specimens of medium- and large-bodied individuals (estimated up to ,8.5 metres long) represent allosauroids. Tyrannosauroids are represented by elements indicating medium body sizes (,3 metres long), likely including the holotype femur of Timimus hermani, and a single cervical vertebra represents a juvenile spinosaurid. Single specimens representing medium- and small-bodied theropods may be referrable to Ceratosauria, Ornithomimosauria, a basal coelurosaur, and at least three taxa within Maniraptora. Thus, nine theropod taxa may have been present. Alternatively, four distinct dorsal vertebrae indicate a minimum of four taxa. However, because most taxa are known from single bones, it is likely that small-bodied theropod diversity remains underestimated. The high abundance of allosauroids and basal coelurosaurs (including tyrannosauroids and possibly ornithomimosaurs), and the relative rarity of ceratosaurs, is strikingly dissimilar to penecontemporaneous dinosaur faunas of Africa and South America, which represent an arid, lower-latitude biome. Similarities between dinosaur faunas of Victoria and the northern continents concern the proportional representatation of higher clades, and may result from the prevailing temperate–polar climate of Australia, especially at high latitudes in Victoria, which is similar to the predominant warm–temperate climate of Laurasia, but distinct from the arid climate zone that covered extensive areas of Gondwana. Most dinosaur groups probably attained a near-cosmopolitan distribution in the Jurassic, prior to fragmentation of the Pangaean supercontinent, and some aspects of the hallmark ‘Gondwanan’ fauna of South America and Africa may therefore reflect climate-driven provinciality, not vicariant evolution driven by continental fragmentation. However, vicariance may still be detected at lower phylogenetic levels. Citation: Benson RBJ, Rich TH, Vickers-Rich P, Hall M (2012) Theropod Fauna from Southern Australia Indicates High Polar Diversity and Climate-Driven Dinosaur Provinciality. PLoS ONE 7(5): e37122. doi:10.1371/journal.pone.0037122 Editor: Andrew A. Farke, Raymond M. Alf Museum of Paleontology, United States of America Received February 11, 2012; Accepted April 13, 2012; Published May 16, 2012 Copyright: ß 2012 Benson et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: Substantial sampling of the Victorian dinosaur fauna would not have occurred without support from National Geographic, Atlas Copco, the Australian Research Council (ARC), Monash University, and Museum Victoria. RBJB’s work on Cretaceous Biogeography and Latitudinal Climatic Variation is supported by a Research Project Grant from the Leverhulme Foundation (RPG-129) awarded to Paul Upchurch. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors’ received funding for field collection of fossils from two commercial sources (Atlas Copco and National Geographic). This does not alter the authors’ adherence to all the PLoS ONE policies on sharing data and materials. * E-mail: [email protected] Introduction South America from East Gondwana (Australia+Antarctica) in the Late Cretaceous, 80 Mya [11–12]. Alternatively, the southern Early Cretaceous (145.5–99.6 million years ago [Mya]; [1]) continents may have all separated within a short interval ,80 Mya tetrapod faunas are of significant interest. Not only do they provide [9]. Both hypotheses suggest enduring physical contact between snapshots of ancient ecosystems and document the early evolution South America, Antarctica and Australia. Thus, prior expectation is of key components of the modern biota (e.g., [2–5]), they also that Early Cretaceous tetrapod faunas of Australia should immediately postdate the Late Jurassic fragmentation of the demonstrate biogeographic links with those of Antarctica and Pangean supercontinent (e.g., [6]). Thus, the topology of evolu- South America. Evidence for such links may include both the tionary branching events among Early Cretaceous tetrapods may representation of higher taxonomic groups, and the phylogenetic constrain our understanding of continental fragmentation, its relationships of individual taxa within those groups. interaction with the evolutionary history of major taxonomic Compared with those of most other continents, the Australian groups, and its role in determining the geographic distribution of dinosaur fauna is poorly-known [13]. Thus, it has been neglected organisms (via cladistic biogeography; e.g., [7–9]). Gondwana, a in most discussions of Cretaceous biogeography (but see [14–17]). landmass made up of the present day southern continents and India, However, recent interpretations of the Australian dinosaur fossils became separated from Laurasia (the northern continents) during have been contentious [15–22]. Debate focuses on the inference of the Jurassic (e.g., [10]). Timing of the subsequent breakup of a ‘Gondwanan’ fauna [23–24], sharing higher taxonomic groups Gondwana is not confidently understood (reviewed by [9]). Africa with South America, Africa, Madagascar and India (recently may have separated early, 140–120 Mya, followed by separation of synthesized by [15]). Alternatively, Australian dinosaur faunas PLoS ONE | www.plosone.org 1 May 2012 | Volume 7 | Issue 5 | e37122 Australian Theropod Dinosaurs may include some classically ‘Laurasian’ clades [16–18,20], abundant collection of teeth from the Flat Rocks Site reflecting near-cosmopolitanism of high-level taxonomic groups (38u39940620S, 145u40952630E). Non-dental cranial elements in the Early Cretaceous (e.g., [16–17,20–21,25–27]). Early are rare, and only a few have been recovered (the surangular Cretaceous cosmopolitanism is congruent with the Jurassic described by [43], and an unidentified frontal), and these are not origination of many groups, known directly from stratigraphic discussed here because we have little to add in terms of occurrences, and inferred from phylogenetic ghost lineages [28– identification. One aim of the present study is to review the 31]. Under this model, clear differences in the higher-taxonomic disputed affinities of the specimens mentioned above, accompa- composition of well-known Late Cretaceous Laurasian and nied by comprehensive figures and detailed comparisons. In Gondwanan dinosaur faunas [24] arose not via continent-level addition, many specimens are described here for the first time, and vicariance, but by differential patterns of regional extinction this new data is highly significant for understanding the global [27,32–33], and local environmental preferences (e.g., in sauro- biogeographic context of the Australian dinosaur fauna. Frag- pods [34–35]). If this is correct, then the signature of North-South mentary Early Cretaceous dinosaur remains, including rare vicariance in Early Cretaceous faunas may be detectable only at theropod elements, are also known from South Australia [46– lower phylogenetic levels; within, but not among, higher clades of 49] and New South Wales (e.g., [46,50–51]), and more complete Jurassic origin. This global biogeographic model does not dispute dinosaur remains from Queensland (e.g., [52–54]), including the the existence of regional faunas with highly-characteristic taxo- holotypic partial skeleton of the allosauroid theropod Australovenator nomic compositions, such as those of Patagonia [14,23] and North wintonensis [45]. We provide only a few comments on this Africa [36–40]. It does suggest, however, that continental additional material. Reviews of Australian dinosaur discoveries fragmentation provides only a partial explanation. of all ages were provided by Rich & Vickers-Rich [55–56] and The biogeographic debate surrounding high-level taxonomic Weishampel et al. [13]. assignment of Australian dinosaur fossils (to either ‘Gondwanan endemics’ or elements of a more global fauna) is underpinned by Institutional abbreviations the disputed identifications of many specimens. Debate centres AMNH, American Museum of Natural History, New York, around specimens from the Early Cretaceous (Aptian–Albian) of USA; IGM, Mongolian Academy of Sciences, Ulaan Baatar, Victoria, most of which are discovered as isolated elements. For Mongolia; IVPP, Institute of Vertebrate Paleontology and Paleo- example, the holotypic ulna of Serendipaceratops arthucclarkei, and anthropology, Beijing, China; MACN, Museo Nacional, Buenos other material, was described as a Protoceratops-like basal ceratop-
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