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A New Clade of Archaic Large-Bodied Predatory Dinosaurs (Theropoda: Allosauroidea) That Survived to the Latest Mesozoic

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A New Clade of Archaic Large-Bodied Predatory Dinosaurs (Theropoda: Allosauroidea) That Survived to the Latest Mesozoic Naturwissenschaften (2010) 97:71–78 DOI 10.1007/s00114-009-0614-x ORIGINAL PAPER A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic Roger B. J. Benson & Matthew T. Carrano & Stephen L. Brusatte Received: 26 August 2009 /Revised: 27 September 2009 /Accepted: 29 September 2009 /Published online: 14 October 2009 # Springer-Verlag 2009 Abstract Non-avian theropod dinosaurs attained large Neovenatoridae includes a derived group (Megaraptora, body sizes, monopolising terrestrial apex predator niches new clade) that developed long, raptorial forelimbs, in the Jurassic–Cretaceous. From the Middle Jurassic cursorial hind limbs, appendicular pneumaticity and small onwards, Allosauroidea and Megalosauroidea comprised size, features acquired convergently in bird-line theropods. almost all large-bodied predators for 85 million years. Neovenatorids thus occupied a 14-fold adult size range Despite their enormous success, however, they are usually from 175 kg (Fukuiraptor) to approximately 2,500 kg considered absent from terminal Cretaceous ecosystems, (Chilantaisaurus). Recognition of this major allosauroid replaced by tyrannosaurids and abelisaurids. We demon- radiation has implications for Gondwanan paleobiogeog- strate that the problematic allosauroids Aerosteon, Austral- raphy: The distribution of early Cretaceous allosauroids ovenator, Fukuiraptor and Neovenator form a previously does not strongly support the vicariant hypothesis of unrecognised but ecologically diverse and globally distrib- southern dinosaur evolution or any particular continental uted clade (Neovenatoridae, new clade) with the hitherto breakup sequence or dispersal scenario. Instead, clades enigmatic theropods Chilantaisaurus, Megaraptor and the were nearly cosmopolitan in their early history, and later Maastrichtian Orkoraptor. This refutes the notion that distributions are explained by sampling failure or local allosauroid extinction pre-dated the end of the Mesozoic. extinction. Electronic supplementary material The online version of this article Keywords Neovenatoridae . Megaraptora . Cretaceous . (doi:10.1007/s00114-009-0614-x) contains supplementary material, Gondwanan biogeography. Dinosaur evolution which is available to authorized users. R. B. J. Benson (*) Department of Earth Sciences, University of Cambridge, Introduction Downing Street, Cambridge CB2 3EQ, UK e-mail: [email protected] Theropod dinosaurs have been an ecologically diverse component of terrestrial ecosystems since their Late M. T. Carrano Triassic origins. Having given rise to birds, they are the Department of Paleobiology, Smithsonian Institution, P.O. Box 37012, MRC 121, most diverse clade of extant amniotes. Non-avian thero- Washington, DC 20013-7012, USA pods, including the characteristic giant predators of the Mesozoic, became extinct at the end of the Cretaceous, 65 S. L. Brusatte million years ago. Prior to this, for 160 million years, Division of Paleontology, American Museum of Natural History, Central Park West at 79th Street, theropods monopolised the role of apex predator in New York, NY 10024, USA terrestrial ecosystems. They produced numerous distinct lineages of large terrestrial predators among ceratosaurs, S. L. Brusatte megalosauroids, allosauroids and tyrannosaurids. Department of Earth and Environmental Sciences, Columbia University, The basal tetanuran clades Allosauroidea and Megalo- New York, NY, USA sauroidea appeared by the Middle Jurassic and were soon 72 Naturwissenschaften (2010) 97:71–78 represented by large-bodied taxa such as Shidaisaurus and several previously identified autapomorphies of Neovenator Megalosaurus (Wu et al. 2009;Benson2009). They are identified here as synapomorphies of a more inclusive yielded the first giant taxa (>2 tonnes) in theropod history clade. shortly thereafter (the megalosauroid Torvosaurus and the Aerosteon, from the Anacleto Formation (Santonian, allosauroid Saurophaganax; Bakker et al. 1992). These two Late Cretaceous) of Argentina, is represented by the clades are central to understanding Middle Jurassic–early holotype partial skeleton (MCNA-PV-3137; see “Electronic Late Cretaceous dinosaurian ecosystems, in which they supplementary material” for institutional abbreviations) and comprised almost all large predators over a span of a partial hind limb (MCNA-PV-3139; MCNA-PV-3138 is approximately 85 million years. Currently, the youngest considered Tetanurae indet; “Electronic supplementary definite allosauroids are from the Turonian of China material”). Preserved cranial bones do not overlap with (Brusatte et al. 2009) and Santonian of Argentina (Sereno material of Neovenator, but the scapulocoracoid, pelvis and et al. 2008). Despite their enormous success, however, parts of the axial column and hind limb are represented. A basal tetanurans are hitherto unknown from the final 20 preliminary report proposed carcharodontosaurid affinities million years of the Mesozoic. Instead, the iconic tyranno- (Alcober et al. 1998), but more recent work concluded that sauroids, a lineage of previously small-bodied coelurosau- it was an allosauroid of uncertain affinities (Sereno et al. rian theropods, achieved masses of several tonnes in 2008). Laurasia (e.g. Erickson et al. 2004). In Gondwana, Aerosteon has never been included in a phylogenetic abelisaurid neoceratosaurs diversified to occupy the apex analysis but shares several features with carcharodontosaur- predator niche (e.g. Sampson and Krause 2007; Carrano ids and Neovenator: double cervical anterior pneumatic and Sampson 2008). foramina, pneumatic foramina in all dorsal vertebrae, Here, we present evidence that six previously enigmatic transversely narrow, sheet-like dorsal hyposphenes theropod taxa—Aerosteon, Australovenator, Chilantaisau- (Fig 1b) and a ventral longitudinal ridge on the proximal rus, Fukuiraptor, Megaraptor and Orkoraptor—form a caudal vertebrae. Aerosteon also shares several features monophyletic clade with the unequivocal allosauroid Neo- with Neovenator and other taxa discussed here (see venator. The Maastrichtian age of Orkoraptor provides “Electronic supplementary material”). Notably, although a conclusive evidence that allosauroids survived until the end pneumatic ilium was originally proposed as unique among of the Cretaceous and were not extinct, as previously non-maniraptoran theropods (Sereno et al. 2008), the thought. Derived members of this new clade exhibit a broken ilia of Neovenator also contain internal pneumatic unique body plan among otherwise conservative basal spaces (Brusatte et al. 2008). tetanurans, highlighted by raptorial forelimbs and cursorial Megaraptor was originally described on the basis of a hind limbs that converge on the anatomy of bird-like fragmentary specimen from the Río Neuquén Formation theropods. In sum, the identification of this new clade, (Turonian–Coniacian) of Neuquén, Argentina, which was composed primarily of taxa that were only recently interpreted as a coelurosaur (Novas 1998), due to the discovered and described (since 1996; other than Chilan- slender proportions of metatarsal III and the supposedly taisaurus), reveals a previously unrecognised major group dromaeosaurid-like pedal ungual. A second specimen, of basal theropods that were an important component of comprising a more complete forelimb, vertebrae, a scap- Mesozoic terrestrial ecosystems. ulocoracoid and a partial pubis from the Portezuelo Formation (Turonian–Santonian), demonstrated that the ungual belonged to manual digit I (Calvo et al. 2004), Comparisons nullifying the proposed similarity with dromaeosaurids. In a global phylogenetic analysis of theropods, Smith et Basic comparisons and synapomorphies are outlined and al. (2007) recovered Megaraptor as a carcharodontosaurid. summarised below (and see Fig. 1). A comprehensive However, Smith et al. (2008) later noted that Megaraptor discussion of the anatomy and taxonomy of the focal taxa shared various features of the forelimb with spinosaurids and a list of specimens examined are included in the and recovered Megaraptor as a megalosauroid (=spinosau- “Electronic supplementary material”. roid). Many of the proposed synapomorphies are problem- Neovenator is known by three specimens from the atic and are interpreted differently here (see “Electronic Wessex Formation (Barremian, Lower Cretaceous) of the supplementary material”). Notably, although the first Isle of Wight, UK. These represent most of the skeleton manual ungual of Megaraptor is large, as in some except the forelimbs and posterior two thirds of the skull megalosauroids (e.g. Charig and Milner 1997), its detailed (Hutt et al. 1996). Numerous skeletal features support a morphology is distinctive. In Megaraptor, the ungual is closer relationship to carcharodontosaurids than to Allosau- transversely narrow, with a ratio of proximal height to rus (Naish et al. 2001; Brusatte et al. 2008). Importantly, width of 2.75 (Fig. 1g–h). This is substantially higher than Naturwissenschaften (2010) 97:71–78 73 Fig. 1 Neovenatorid anatomy shown by Aerosteon (a–b, d–e MCNA- g–h right phalanx I; i right tibia; j right tibia (reversed); and k left PV-3137; k MCNA-PV-3139), Megaraptor (c, f, g–h MUCPv 341), tibia. acdl anterior centrodiapophyseal lamina, asp ascending process, Neovenator (i MIWG 6348) and Chilantaisaurus (j IVPP V.2884) in cnc cnemial crest, fos fossa, hp hyposphene, lpr anterior process of ventral (a, f, h), posterior (b), right lateral (c), medial (d, g), lateral (e) lateral
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