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Cretaceous of Australia the Origin of Modern Crocodyliforms

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Cretaceous of Australia the Origin of Modern Crocodyliforms View metadata, citation and similar papers atDownloaded core.ac.uk from rspb.royalsocietypublishing.org on November 23, 2010 brought to you by CORE provided by RERO DOC Digital Library The origin of modern crocodyliforms: new evidence from the Cretaceous of Australia Steven W Salisbury, Ralph E Molnar, Eberhard Frey and Paul M.A Willis Proc. R. Soc. B 2006 273, 2439-2448 doi: 10.1098/rspb.2006.3613 Supplementary data "Data Supplement" http://rspb.royalsocietypublishing.org/content/suppl/2009/03/12/273.1600.2439.DC1.ht ml References This article cites 37 articles, 4 of which can be accessed free http://rspb.royalsocietypublishing.org/content/273/1600/2439.full.html#ref-list-1 Article cited in: http://rspb.royalsocietypublishing.org/content/273/1600/2439.full.html#related-urls Receive free email alerts when new articles cite this article - sign up in the box at the top Email alerting service right-hand corner of the article or click here To subscribe to Proc. R. Soc. B go to: http://rspb.royalsocietypublishing.org/subscriptions This journal is © 2006 The Royal Society Downloaded from rspb.royalsocietypublishing.org on November 23, 2010 Proc. R. Soc. B (2006) 273, 2439–2448 doi:10.1098/rspb.2006.3613 Published online 14 June 2006 The origin of modern crocodyliforms: new evidence from the Cretaceous of Australia Steven W. Salisbury1,2,*, Ralph E. Molnar3, Eberhard Frey4 and Paul M. A. Willis5 1School of Integrative Biology, The University of Queensland, Brisbane, Qld 4072, Australia 2Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA 15213-4080, USA 3Museum of Northern Arizona, 3101 North Fort Valley Road, Flagstaff, AZ 86001, USA 4Staatliches Museum fu¨r Naturkunde Karlsruhe, Erbprinzenstrasse 13, 76133 Karlsruhe, Germany 5School of Biological Sciences, University of New South Wales, Sydney, NSW 2052, Australia While the crocodyliform lineage extends back over 200 million years (Myr) to the Late Triassic, modern forms—members of Eusuchia—do not appear until the Cretaceous. Eusuchia includes the crown group Crocodylia, which comprises Crocodyloidea, Alligatoroidea and Gavialoidea. Fossils of non-crocodylian eusuchians are currently rare and, in most instances, fragmentary. Consequently, the transition from Neosuchia to Crocodylia has been one of the most poorly understood areas of crocodyliform evolution. Here we describe a new crocodyliform from the mid-Cretaceous (98–95 Myr ago; Albian–Cenomanian) Winton Formation of Queensland, Australia, as the most primitive member of Eusuchia. The anatomical changes associated with the emergence of this taxon indicate a pivotal shift in the feeding and locomotor behaviour of crocodyliforms—a shift that may be linked to the subsequent rapid diversification of Eusuchia 20 Myr later during the Late Cretaceous and Early Tertiary. While Laurasia (in particular North America) is the most likely ancestral area for Crocodylia, the biogeographic events associated with the origin of Eusuchia are more complex. Although the fossil evidence is limited, it now seems likely that at least part of the early history of Eusuchia transpired in Gondwana. Keywords: Crocodyliformes; Eusuchia; Crocodylia; evolution; Cretaceous; Gondwana 1. INTRODUCTION back to the Campanian (ca 80 Myr ago; alligatoroids and Crocodyliforms are the only non-avian archosaurs to have gavialoids) and late Maastrichtian (ca 67–65 Myr ago; persisted through the Mesozoic and Cenozoic into crocodyloids; Brochu 1997, 2001). Fossils of more basal modern times. With their stark appearance, it is often forms, however, such as the advanced neosuchian assumed that they have changed little in 200 million years Bernissartia (Norell & Clark 1990), are much older (Myr) of evolution. Yet the morphological diversity that (Hauterivian–lower Aptian; ca 130–120 Myr ago). crocodyliforms display today represents only a fraction of The transition between advanced neosuchians and that during the Mesozoic when several distinct suborders crocodylians involved subtle but far-reaching changes to existed (Clark 1994). Only one of these suborders, the entire body. Sir Thomas Huxley was the first to Eusuchia, still exists. recognize these changes, regarding them as one of the Eusuchia includes all extant crocodyliforms: members strongest cases yet for Darwin’s then recently published of the crown group Crocodylia (Benton & Clark 1988). theory of evolution, and integrated them into the first Crocodylia comprises Crocodyloidea (which includes formal classification of both fossil and living crocodyli- extant species of Crocodylus and Osteolaemus tetraspis), forms (Huxley 1875). Huxley identified three main phases Alligatoroidea (alligators and caimans) and Gavialoidea in the evolution of crocodyliforms: ‘Parasuchia’, ‘Meso- (the group that includes the Indian gharial, Gavialis suchia’ and Eusuchia. His definition of Eusuchia was apomorphy-based, with referral to the group dependent gangeticus). The position of the Sunda or false gharial, on the combined possession of: (i) a fully developed bony Tomistoma schlegelii, floats between Crocodyloidea palate, where the secondary choanae are fully enclosed (morphological studies: e.g. Norell 1989; Salisbury & mediorostrally by ventral laminae of the pterygoids; and Willis 1996; Brochu 1997, 1999, 2004) and Gavialoidea (ii) procoelous vertebrae, in which there are synovial, (biochemical or molecular studies: e.g. Densmore & semi-spheroidal articulations between adjoining vertebral Owen 1989; Gatesy et al. 2003; Harshman et al. 2003). bodies (Salisbury & Frey 2001). Although these features Fossils that can be referred to each of the three are now known to have evolved independently in other superfamilies of Crocodylia are well known, extending crocodyliforms (Michard et al. 1990; Rogers 2003; Clark et al. 2004), the presence of both features in combination * Author for correspondence ([email protected]). with a sagitally segmented paravertebral shield has thus far proven to be limited to eusuchians and, therefore, The electronic supplementary material is available at http://dx.doi. org/10.1098/rspb.2006.3613 or via http://www.journals.royalsoc.ac. phylogenetically informative (Norell & Clark 1990; uk. Clark & Norell 1992). Received 24 April 2006 2439 q 2006 The Royal Society Accepted 4 May 2006 Downloaded from rspb.royalsocietypublishing.org on November 23, 2010 2440 S. W. Salisbury and others The origin of modern crocodyliforms along with tooth crowns from the left maxilla (figure 4e,f ); QM F34642, a partial, articulated skeleton. (d) Locality and horizon The holotype was found at field locality QM L1021, near the town of Isisford, central-western Queensland, Australia (figure 1). All the preserved portions of the holotype were found in close association, in adjoining portions of a large nodule of fluvial, volcanoclastic sandstone. Most of the holotype and QM F34642 were Figure 1. Mid-Cretaceous (latest Albian–earliest Cenomanian; found during the mid-1990s, with the remaining portions 98–95 Myr ago)palaeogeographic map, (Mollweideprojection, discovered during 2001 and 2003. Other referred speci- latitudeand longitudelines at 308 intervals;after Dettmann et al. mens were found in the same horizon, at sites in the 1992; Scotese 2001). White star indicates fossil locality. vicinity of the type locality during University of Queens- Unfortunately, the transition itself—the point at which land expeditions between 2003 and 2005. neosuchians become eusuchians, and eusuchians in turn The horizon in which the specimens occur is part of the become crocodylians—has been obscure, with the known Winton Formation. This formation spans the transition fossils tending to be incomplete, badly preserved or between the Early Cretaceous and the Late Cretaceous undescribed (Molnar 1980; Clark & Norell 1992; (latest Albian–earliest Cenomanian; Helby et al. Buscalioni et al. 2001). Consequently, the origins of 1987)—atimethatweinformallyrefertoasthe Eusuchia and the subsequent emergence of taxa immedi- ‘mid-Cretaceous’. The Winton Formation has previously ately ancestral to Crocodylia have remained one of the produced the remains of titanosauriform sauropods most poorly understood areas of crocodyliform evolution (Molnar & Salisbury 2005), ceratodont lungfishes (Huene 1933; Benton & Clark 1988; Norell & Clark 1990; (Dettmann et al.1992) and vertebrate microfossils Clark & Norell 1992; Brochu 1999). referable to theropods, thyreophorans, turtles and possible Herein we report on a new crocodyliform from mammaliaforms. Trackways also point to the presence of Australia that provides new evidence on the transition two types of theropod and at least two types of ornithopod from Neosuchia to Crocodylia during the Early Cret- (Thulborn & Wade 1984). aceous. Among the material referred to, the new taxon is a complete skull and an almost complete, fully articulated (e) Differential diagnosis skeleton—the earliest such example from a definitive basal Isisfordia differs from other crocodyliforms in the following eusuchian and the most complete crocodyliform fossil unique combination of traits (autapomorphies marked with from Australia. an ‘a’): broad exposure of the exoccipital within the supratemporal foramen rostral to the rostral aperture of the posttemporal canal (a); maximum diameter of the caudal 2. MATERIAL AND METHODS aperture of the cranioquadrate siphonium approximately The material described here has been accessioned to the one-third the mediolateral width of the foramen magnum, Queensland Museum (QM), Brisbane, Australia. The with
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