The phylogeny of Tetanurae (Dinosauria: Theropoda)

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The phylogeny of Tetanurae (Dinosauria: Theropoda) a

b

Matthew T. Carrano , Roger B. J. Benson & Scott D. Sampson

c

a

Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560, USA b

Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge, CB2 3EQ, UK c

Natural History Museum of Utah and Department of Geology & Geophysics, University of Utah, Salt Lake City, UT, 84108, USA Available online: 17 May 2012

To cite this article: Matthew T. Carrano, Roger B. J. Benson & Scott D. Sampson (2012): The phylogeny of Tetanurae (Dinosauria: Theropoda), Journal of Systematic Palaeontology, 10:2, 211-300 To link to this article: http://dx.doi.org/10.1080/14772019.2011.630927

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Journal of Systematic Palaeontology, Vol. 10, Issue 2, June 2012, 211–300

FEATURED ARTICLE

The phylogeny of Tetanurae (Dinosauria: Theropoda) Matthew T. Carranoa∗, Roger B. J. Bensonb and Scott D. Sampsonc a

Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA; bDepartment of Earth Sciences, University of Cambridge, Downing Street, Cambridge, CB2 3EQ, UK; cNatural History Museum of Utah and Department of Geology & Geophysics, University of Utah, Salt Lake City, UT 84108, USA

Downloaded by [Smithsonian Institution Libraries] at 07:12 17 May 2012

(Received 9 November 2010; accepted 19 June 2011; printed 16 May 2012) Tetanuran theropods represent the majority of Mesozoic predatory dinosaur diversity and the lineage leading to extant Aves. Thus their history is relevant to understanding the evolution of dinosaur diversity, Mesozoic terrestrial ecosystems, and modern birds. Previously, the fragmentary and poorly sampled fossil record of basal (non-coelurosaur) tetanurans led to uncertainties regarding their basic interrelationships. This in turn prevented determining the relationships of many incompletely known taxa that nonetheless document a global radiation spanning more than 120 million years. We undertook an exhaustive examination of all basal tetanurans and all existing character data, taking advantage of recent discoveries and adding new morphological, temporal and geographic data. Our cladistic analysis of 61 taxa achieved significantly improved phylogenetic resolution. These results position several ‘stem’ taxa basal to a succession of monophyletic clades (Megalosauroidea, Allosauroidea and Coelurosauria). Megalosauroids include nearly 20 taxa arrayed amongst a basalmost clade (Piatnitzkysauridae, fam. nov.) and the sister taxa Spinosauridae and Megalosauridae; the latter includes two subfamilies, Megalosaurinae and Afrovenatorinae subfam. nov. Allosauroidea contains a diverse Metriacanthosauridae (= Sinraptoridae), Neovenatoridae, Carcharodontosauridae and a reduced Allosauridae. Finally, we assessed more than 40 fragmentary forms and hundreds of additional reported tetanuran occurrences. Tetanuran evolution was characterized by repeated acquisitions of giant body size and at least two general skull forms, but few variations in locomotor morphology. Despite parallel diversification of multiple lineages, there is evidence for a succession of ‘dominant’ clades. Tetanurae first appeared by the Early Jurassic and was globally distributed by the Middle Jurassic. Several major clades appeared prior to the breakup of Pangaea; as such their absence in specific regions, and at later times, must be due to poor sampling, dispersal failure and/or regional extinction. Finally, we outline a general perspective on Mesozoic terrestrial biogeography that should apply to most clades that appeared before the Late Jurassic. Keywords: evolution; systematics; Mesozoic; taxonomy; morphology, biogeography

Introduction Theropod biology, evolution and phylogeny have received disproportionate attention within the study of dinosaur palaeontology. Currently, non-avian theropod taxa comprise more than 40% of all named dinosaur species, more than any other major clade (Weishampel et al. 1990, 2004a). It is tempting to interpret this as a reflection of an unusually diverse dinosaur clade, but this perceived diversity is heavily mitigated by certain anthropogenic biases. Certainly the inherently dramatic predatory qualities of these organisms have lent them a special focus, but significant research efforts have also been directed at resolving the problems surrounding bird origins (e.g. Gauthier 1986; Turner et al. 2007). As a result, theropods are probably both oversampled and overstudied relative to other dinosaur groups. In spite of this, many individual theropod taxa are as poorly studied as most other dinosaurs. Because research ∗

Corresponding author. Email: [email protected]

ISSN 1477-2019 print / 1478-0941 online C 2012 The Natural History Museum Copyright http://dx.doi.org/10.1080/14772019.2011.630927 http://www.tandfonline.com

has tended to focus on the especially impressive (e.g. Tyrannosaurus) and the nearly avian (e.g. Dromaeosauridae) amongst the Theropoda, much less has been learnt about the 100+ more primitive forms. Indeed, those aspects most fundamental to understanding the evolution of basal theropods – their phylogenetic interrelationships – have only recently received much specific attention (e.g. Sereno et al. 1996, 1998; Holtz 2000, 2004; Rauhut 2003; Carrano & Sampson 2008; Benson 2010a). Most previous studies of basal tetanurans have focused on placing individual, usually new, taxa into a phylogenetic context (e.g. Sereno et al. 1994, 1996) and tend to include more complete forms in the hope of achieving the greatest resolution. By contrast, other works have attempted to resolve relationships amongst theropods as a whole (e.g. Holtz 2000; Rauhut 2003). In general (see below) these have produced more limited resolution, often conflicting in significant details with one another. This probably arises from a paucity of characters constructed specifically to


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resolve basal tetanuran relationships (rather than particular ingroups of closely related taxa), which itself results from the historical lack of focused study. Several years ago, two of us (MTC and SDS) undertook a major project to address these difficulties. We began with a comprehensive review of theropod interrelationships, focusing on all taxa more primitive than Coelurosauria. This work has involved first-hand examination of hundreds of theropod specimens, critical review of nearly 700 existing characters from all published studies, and inclusion of many taxa and characters never previously incorporated into any phylogenetic analyses. RBJB independently undertook a review focused on non-coelurosaurian (‘basal’) tetanurans and also reviewed existing taxa and character descriptions. The present work represents a combination of our efforts. This paper is the sixth documenting our phylogenetic results. The first (Carrano et al. 2002) provided evidence that Ceratosauria was a lineage independent of Coelophysoidea and more closely allied with Tetanurae. The second (Carrano et al. 2005) produced a phylogeny of Coelophysoidea along with a redescription of the small theropod Segisaurus. The third (Carrano & Sampson 2008) focused on the ingroup relationships of Ceratosauria. Benson (2010a) focused on resolving the relationships of Middle Jurassic theropods, including the problematic European ‘megalosaurs’, and we recently recognized a distinct lineage of Cretaceous allosauroids (Benson et al. 2010). These works complement several descriptive studies of individual theropod taxa, including the ceratosaurs Majungasaurus (Carrano 2007; Sampson & Witmer 2007) and Masiakasaurus (Sampson et al. 2001; Carrano et al. 2002; Carrano et al. 2011), the basal tetanuran Monolophosaurus (Brusatte et al. 2010a; Zhao et al. 2010), the megalosaurs Duriavenator, Magnosaurus and Megalosaurus (Benson 2008a, 2010a, b), the carcharodontosaurians Chilantaisaurus, Kelmayisaurus, Neovenator, and Shaochilong (Benson & Xu 2008; Brusatte et al. 2008, 2010b, 2011), and the coelurosaur Stokesosaurus (Benson 2008b). Together these have provided new anatomical and character data for the present study. Here we present a systematic analysis with the goal of producing a well resolved, highly documented and comprehensive phylogeny of all basal (= non-coelurosaurian) tetanurans. Several non-tetanurans and coelurosaurs are included as reference taxa to define the basic interrelationships of the major theropod clades; ceratosaurs are thus included but at a reduced level of sampling. Although we employ species level sampling and avoid using suprageneric groups, we have excluded a number of fragmentary forms in an attempt to achieve some measure of resolution. When possible, we have used criteria for Safe Taxonomic Reduction, omitting only those highly incomplete taxa whose character combinations are not unique (Wilkinson 1995). Additionally we employ Adams consensus methods (Adams 1972) to identify ‘wildcard’

taxa a posteriori. This is a manual implementation of the strict reduced consensus method (e.g. Wilkinson 2003).

Historical background Constituency and placement of Tetanurae Although the term is commonly used today in dinosaur studies, Tetanurae was only recently recognized and named (Gauthier 1986). Gauthier’s study is an important benchmark for theropod systematics as it represents the earliest comprehensive cladistic treatment of Theropoda and the source of much of the basic topology on which current studies are still constructed. That said, the taxa now called tetanurans include the earliest named dinosaur (Megalosaurus Buckland, 1824), and a wide array of forms discovered since that span nearly the entire temporal and geographic range of Dinosauria. As such, these taxa have been of interest to theropod and dinosaur workers since the 19th century, even though they have rarely been recognized as belonging to a single evolutionary lineage. Indeed, within the ranks of Tetanurae are found some of the most problematic taxa in all Dinosauria. Megalosaurus has achieved an almost mythic status among these. A fragmentarily known form from the Middle Jurassic of England, all known topotypic specimens are dissociated and were deposited allochthonously in marginal marine sediments. Its early discovery and description (Buckland 1824) predate both the term Dinosauria and any standardization of palaeontological reporting, thereby contributing to the confusion surrounding its identity and relationships. Unfortunately, as a hallmark theropod, Megalosaurus served as a taxonomic ‘attractor’ for decades, resulting in the unstable, often incorrect, and usually unjustified placement of dozens of referred species within the genus. This is discussed in further detail below. For a century after the description of Megalosaurus, most large carnivorous dinosaurs were grouped into the family Megalosauridae within the order Theropoda (e.g. Baur 1891; Hay 1902; Romer 1956; Walker 1964; Fig. 1). At times distinct European and North American lineages were recognized, with the latter often referred to as Dryptosauridae (Marsh 1895). Occasionally, workers separated individual taxa into discrete families if their morphology appeared aberrant (e.g. Labrosaurus, Labrosauridae; Ceratosaurus, Ceratosauridae; Spinosaurus, Spinosauridae), but Megalosauridae remained the only multi-taxic group of large theropods employed consistently during this time. In addition, nearly all researchers arrayed the families of carnivorous dinosaurs serially within Theropoda (or Saurischia), with only a few forms segregated into taxa of higher rank (e.g. Ceratosauria, Coeluria, Compsognatha).


The phylogeny of Tetanurae

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Huxley 1870

Williston 1878

Marsh 1882

Lydekker 1888

Baur 1891

Dinosauria Iguanodontidae Megalosauridae Euskelesaurus Laelaps Megalosaurus Palaeosaurus Poekilopleuron Teratosaurus Scelidosauridae

Dinosauria Ornithoscelida Allosaurus Compsognathus Creosaurus Dryptosaurus Hypsilophodon Iguanodon Laosaurus Nanosaurus Poekilopleuron Sauropoda

Theropoda Amphisauridae Compsognatha Dryptosauridae Dryptosaurus Labrosauridae Labrosaurus Megalosauridae Allosaurus Coelosaurus Creosaurus Megalosaurus Zanclodontidae

Theropoda Anchisauridae Coeluridae Compsognathidae Megalosauridae Aristosuchus Bothriospondylus Dryptosaurus Megalosaurus Zanclodon

Megalosauria Anchisauridae Coeluridae Compsognathidae Dryptosauridae Dryptosaurus Megalosauridae Ceratosaurus Megalosaurus Zanclodontidae

Marsh 1895, 1896

Hay 1902

Huene 1909

Theropoda Ceratosauria Compsognatha Hallopoda Coeluria Anchisauridae Dryptosauridae Allosaurus Coelosaurus Creosaurus Dryptosaurus Labrosauridae Labrosaurus Megalosauridae Megalosaurus Plateosauridae

Theropoda Anchisauridae Ceratosauridae Coeluridae Hallopidae Megalosauridae Allosaurus Antrodemus Coelosaurus Creosaurus Deinodon Dryptosaurus Megalosaurus Palaeoctonus Suchoprion Troodon Zapsalis Zatomus Ornithomimidae

Saurischia Ceratosaurus Coeluridae Compsognathidae Megalosauridae Allosaurus Antrodemus Creosaurus Labrosaurus Megalosaurus Plateosauridae Sauropoda Sellosauridae Thecodontosauridae Zanclodontidae

Theropoda Anchisauridae Coeluridae Compsognathidae Labrosauridae Labrosaurus Megalosauridae Allosaurus Antrodemus Coelosaurus Creosaurus Dryptosaurus Laelaps Megalosaurus

Plateosauridae Euskelosaurus Gresslyosaurus Pachysaurus Plateosaurus Poikilopleuron Sellosaurus Teratosaurus Zanclodon

Huene 1914d

Gilmore 1920

Huene 1923b

Huene 1926a

Huene 1926b

Theropoda Ceratosauridae Coeluridae Megalosauridae Antrodemus Deinodon Dryptosaurus Labrosaurus Tyrannosaurus Ornithomimidae

Coelurosauria Carnosauria Megalosauridae Altispinax Antrodemus Megalosaurus Streptospondylus Cladeiodon Dryptosaurus Erectopus Spinosaurus Teratosaurus Zanclodon

Coelurosauria Pachypodosauria Carnosauria Megalosauridae Altispinax Erectopus Megalosaurus Prosauropoda Sauropoda

Coelurosauria Carnosauria Megalosauridae Antrodemus Megalosaurus Spinosauridae Spinosaurus

Coelurosauria Pachypodosauria Ammosauridae Massospondylidae Megalosauridae Plateosauridae Sauropoda Sellosauridae Zanclodontidae

Huene 1929

Huene 1959

Coelurosauria Pachypodosauria Carnosauria Dinodontidae Megalosauridae Prosauropoda Sauropoda

Coelurosauria Carnosauria Allosauridae Dinodontidae Teratosauridae

Zittel 1911

Romer 1956 Theropoda Coelurosauria Carnosauria Megalosauridae Acrocanthosaurus Antrodemus Bahariasaurus Carcharodontosaurus Ceratosaurus Chienkosaurus Dryptosauroides Dryptosaurus

Embasaurus Erectopus Macrodontophion Megalosaurus Orthogoniosaurus Sarcosaurus Spinosaurus Szechuanosaurus Palaeosauridae Teratosauridae Tyrannosauridae

Figure 1. Historical classification schemes that have included theropods which are now classified as tetanurans, 1870–1956. Taxa recovered as non-coelurosaurian tetanurans in the present study are indicated in bold.


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In the early 20th century, Huene produced several reviews of the large theropods then known, resulting in a series of increasingly refined classifications. He was primarily concerned with the overall systematics of Saurischia, a topic he worked on for nearly 50 years (e.g. Huene 1914a, b, c, d, 1920, 1923, 1926a, b, 1932, 1956). He described many new taxa, some of which are still considered valid, but his most significant achievement was the creation of an underlying systematic scheme for predatory dinosaurs that effectively replaced the serial arrangement of previous decades. Initially Huene (1914a, b, c, d) arranged the small, lightly built forms into the infraorder Coelurosauria and put the larger bodied taxa into the infraorder Pachypodosauria, although still within the family Megalosauridae (Pachypodosauria included many forms now considered to be prosauropods and/or basal sauropodomorphs). Later, however, Huene (1920) transferred the large, strictly carnivorous taxa from the Pachypodosauria to a new infraorder, Carnosauria. The remaining pachypodosaurs were allied with sauropods in the Sauropodomorpha. Carnosauria (Huene 1923, 1926a, b) then included all known large bodied predators other than Ceratosaurus and tyrannosaurids (Albertosaurus, Gorgosaurus, Tyrannosaurus). The former was retained within Coelurosauria on the line leading from Coelophysis; the latter were considered to be closely related to ornithomimids. Although both were placed in Carnosauria by many subsequent workers, the arrangement posited by Huene (1926a, b) is remarkably close to our current perception of theropod relationships. A few workers presented somewhat different views during this time. Unlike Huene, Matthew (1915) retained large theropods in Theropoda, apart from Coelurosauria. Likewise, Nopcsa (1923) separated the large theropods from pachypodosaurs into a separate Megalosauria. Although he retained a distinction between them and smaller theropods (Coelurosauria), the three groups were of equivalent rank within Saurischia, alongside Sauropoda. Neither of these had the lasting influence of Huene’s schemes, nor did any of the other variations produced by different workers. But nearly every study emphasized the phylogenetic differences between large and small theropods, and placed basal tetanurans among the former. The size-based distinction between carnosaurs and coelurosaurs rooted in Huene’s classification had a long lasting effect on theropod systematics; for more than half a century, few workers presented serious challenges to this organizational scheme. Even the later compilations of Romer (1956, 1966) and Steel (1970) retained this basic structure (Fig. 2). Aside from acceptance of Spinosauridae and Allosauridae as separate families (e.g. Colbert & Russell 1969; Carroll 1988), little changed for nearly 60 years. Among traditional studies, only Kurzanov (1989)

erected new families of large theropods (Streptospondylidae and Torvosauridae) within the Carnosauria. Walker’s (1964) study separated megalosaurids and spinosaurids into different superfamilies, and thereby provided more structure than previous classifications, but still maintained both within the Carnosauria. Not until Gauthier (1986) fundamentally reorganized theropod interrelationships were any larger changes established. And although Gauthier retained the terms Carnosauria and Coelurosauria, as well as the duality of their relationship, his redefinitions effectively replaced earlier size based concepts. Modern phylogenetic analyses generally agree on a monophyletic Tetanurae that includes a series of generally large bodied basal forms that lie outside a monophyletic Coelurosauria (Figs 3–5). Tyrannosauridae is now universally included within Coelurosauria (Novas 1991a; Holtz 1994a), whereas ceratosaurs and coelophysoids are basal to Tetanurae. Although some workers prefer the term Carnosauria for allosaurs and their closest relatives (e.g. Holtz 2000; also termed Allosauroidea; Sereno et al. 1994), the term does not include all basal forms or all large bodied theropods. However, its use as the name for a possible clade comprising Allosauroidea and Megalosauroidea (Spinosauroidea) as employed by Rauhut (2003) is faithful to Huene’s (1920, 1923a, b, 1926a, b) original definition.

Ingroup relationships of Tetanurae The interrelationships of individual tetanuran taxa have been even more problematic than their basic organization. The prevalence of serially arranged families within most early works (e.g. Marsh 1882, 1895; Hay 1902) often meant that within-family relationships were paid less attention. Alternatively, Huene (1923a, b, 1926a, b) and others tended to describe specific evolutionary lineages of theropods, in which particular taxa were ancestral to others. There is some analogy to the sister-taxon concept of modern cladistic discussion only in the sense that particular lowerlevel affinities are implied between individual taxa. Initial cladistic studies of tetanuran theropods (Paul 1984; Holtz 1994a, 2000; Charig & Milner 1997) generally supported a primitive grade of ‘megalosaurs’ (Eustreptospondylus, Megalosaurus, Torvosaurus) that were arranged as serial outgroups to a small clade of allosaurs (minimally Allosaurus and Acrocanthosaurus), followed by the Coelurosauria. The consensus was that ‘megalosaurs’ represented basal tetanurans, and thus taxa more derived than ‘ceratosaurs,’ but lacked synapomorphies that might support their monophyly (Fig. 3). Subsequently, most workers have recognized that many of these basal tetanurans formed a true clade, usually termed Spinosauroidea (originally Torvosauroidea [Sereno et al. 1994; Olshevsky, 1995] but correctly Megalosauroidea [Holtz et al. 2004; Benson 2010a]). Current disagreements centre on whether this clade is basal to the Allosauroidea



215

Walker 1964

Colbert 1964

Romer 1966

Colbert & Russell 1969

Theropoda Coelurosauria Carnosauria Megalosauroidea Megalosauridae Antrodemus Bahariasaurus Carcharodontosaurus Dryptosaurus Eustreptospondylus Indosaurus Megalosaurus Metriacanthosaurus Tyrannosauroidea Ornithosuchidae Spinosauridae Acrocanthosaurus Altispinax Spinosaurus Tyrannosauridae

Theropoda Coelurosauria Carnosauria Megalosauridae Tyrannosauridae

Theropoda Coelurosauria Carnosauria Megalosauridae Allosaurus Bahariasaurus Carcharodontosaurus Ceratosaurus Chienkosaurus Chilantaisaurus Dryptosauroides Dryptosaurus Embasaurus Erectopus Eustreptospondylus Inosaurus Macrodontophion Megalosaurus Metriacanthosaurus Proceratosaurus Sarcosaurus Ornithosuchidae Poposauridae Spinosauridae Acrocanthosaurus Altispinax Spinosaurus Tyrannosauridae

Theropoda Coelurosauria Carnosauria Allosauridae Acrocanthosaurus Allosaurus Megalosauridae Ceratosaurus Eustreptospondylus Spinosauridae Tyrannosauridae

Swinton 1970

Barsbold 1983

Russell 1984

Welles 1984

Theropoda Coelurosauria Carnosauria Dinodontidae Megalosauridae Acrocanthosaurus Altispinax Antrodemus Ceratosaurus Megalosaurus Metriacanthosaurus Spinosaurus

Theropoda Coelurosauria Carnosauria Allosauridae Ceratosauridae Megalosauridae Tyrannosauridae Spinosauridae

Theropoda Coelurosauria Carnosauria Allosauridae Allosaurus Aublysodontidae Ceratosauridae Dryptosauridae Megalosauridae Acrocanthosaurus Marshosaurus Stokesosaurus Torvosaurus Tyrannosauridae

Theropoda Ceratosauridae Coelophysidae Coeluridae Compsognathidae Dromaeosauridae Halticosauridae Marshosaurus Megalosauridae Allosaurus Coelosaurus Eustreptospondylus Megalosaurus Teratosaurus Procompsognathidae

Carroll 1988 Theropoda Chilantaisaurus Marshosaurus Allosauridae Allosaurus Indosaurus Piatnitzkysaurus Piveteausaurus Yangchuanosaurus Ceratosauridae Coeluridae Compsognathidae Deinocheiridae Dromaeosauridae Dryptosauridae Elmisauridae Megalosauridae Bahariasaurus Carcharodontosaurus Chingkankousaurus Embasaurus Erectopus

Eustreptospondylus Inosaurus Kelmayisaurus Majungasaurus Megalosaurus Metriacanthosaurus Poekilopleuron Szechuanosaurus Torvosaurus Xuanhanosaurus Ornithomimidae Oviraptoridae Podokesauridae Saurornithoididae Shanshanosauridae Spinosauridae Acrocanthosaurus Altispinax Spinosaurus Therizinosauridae Tyrannosauridae

Kurzanov 1989 Carnosauria Abelisauridae Allosauridae Acrocanthosaurus Allosaurus Chilantaisaurus Compsosuchus Marshosaurus Ornithomimoides Piatnitzkysaurus Szechuanosaurus Megalosauridae Gasosaurus Iliosuchus Magnosaurus Megalosaurus Metriacanthosaurus Piveteausaurus Sarcosaurus Yangchuanosaurus

Spinosauridae Altispinax Spinosaurus Streptospondylidae Eustreptospondylus Streptospondylus Torvosauridae Erectopus Poekilopleuron Torvosaurus Tyrannosauridae

Figure 2. Historical classifications, 1956–1989. Taxa recovered as non-coelurosaurian tetanurans in the present study are indicated in bold.

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216 M. T. Carrano et al.

Paul 1984

Novas 1992

Sereno et al., 1994 Paul 1988a

Russell & Dong 1993

Coria & Salgado 1995 Holtz 1994a

Sereno et al., 1996

Figure 3. Results of previous phylogenetic analyses of Tetanurae, 1984–1996. Taxa recovered as non-coelurosaurian tetanurans in the present study are indicated in bold.

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Charig & Milner 1997 Sereno et al., 1998

Allain 2002

+ Coelurosauria clade (usually called Avetheropoda or Neotetanurae; Paul 1988a; Sereno et al. 1994, 1996, 1998; Holtz 2000; Holtz et al. 2004), or instead is the sister taxon to Allosauroidea within a reconsti-

217

Holtz, 2000

Rauhut, 2003

Holtz et al., 2004


tuted Carnosauria (Currie 1995; Rauhut 2003) (Figs 3, 4). The placement of many individual taxa within any of these frameworks also varies. For example, allosauroids

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Co el o Zu ph pa yso y “d sau idea ilo ru CE pho s RA sa Pi TO urs at SA ” n U Co itzk RIA nd ys Irr or aur it ra u Su ato pto s ch r r Ba om ry im o u NM ny s V x Eu P1 s t 86 St rept 07 re os 6 p M tos pon eg po dy a n l To rap dy us rv t lu Du osa or s br ur Af eui us ro llo M ven sau on at ru Al olo or s lo ph s Ne aur osa o u uru Si ven s s nr at a Ac pto or ro r Ty ca ra nt n h Ca not osa rc ita uru Gi har n s ga od n o M oto nto ap sa sa u u u CO sau rus rus EL rus UR OS AU RI A

218 M. T. Carrano et al.

Smith et al., 2007

Benson et al., 2010



The phylogeny of Tetanurae reliably comprise Allosaurus, Acrocanthosaurus, Neovenator, Sinraptoridae and Carcharodontosauridae, but there is disagreement about whether Neovenator and Acrocanthosaurus are allied with allosaurids (Currie & Carpenter 2000; Holtz 2000; Allain 2002; Novas et al. 2005; Smith et al. 2007) or carcharodontosaurids (Sereno et al. 1996; Rauhut 2003; Brusatte & Sereno 2008; Benson 2010a; Benson et al. 2010). Monolophosaurus, Fukuiraptor, Cryolophosaurus and Piatnitzkysaurus have all been placed within Allosauroidea (Bonaparte 1986; Sereno et al. 1994) but each exhibits morphological evidence of other affinities (Zhao & Currie 1994; Rauhut 2003; Holtz et al. 2004; Smith et al. 2007; Benson 2010a; Brusatte et al. 2010a; Zhao et al. 2010). ‘Megalosaurs’ pose an even greater and more complex problem. Many of the taxa that have at one time been referred to Megalosauridae have now been dispersed elsewhere, but a large number of putative megalosaur species remain. These include forms that have also been called torvosaurids (Kurzanov 1989; Sereno et al. 1994, 1996), streptospondylids (Kurzanov 1989) and eustreptospondylids/eustreptospondylines (Paul 1988a; Bakker et al. 1992; Yates 2005), but which are now considered members of Spinosauroidea (Megalosauroidea). Megalosaurus has been suggested to belong here (Britt 1991) but this was only recently supported analytically (Benson, 2010a). Similarities between Torvosaurus, Edmarka and Eustreptospondylus have been noted (Bakker et al. 1992), as well as between these and spinosaurids (Sereno et al. 1994, 1996, 1998). More recent work has placed additional taxa within this group, including Streptospondylus (Allain 2001; Smith et al. 2007), Dubreuillosaurus (Allain 2002, 2005a; Benson 2010a) and Poekilopleuron (Allain & Chure 2002; Holtz et al. 2004). Despite some agreement supporting a diverse spinosauroid (or megalosauroid) clade, there is less consensus regarding the relationships between these taxa, resulting in little consistent support for anything other than a spinosaurid/torvosaurid (megalosaurid) dichotomy. Other taxa remain ambiguously placed. For example, Afrovenator has been allied with both allosauroids (Rauhut 2003) and spinosauroids (Sereno et al. 1994; Allain 2002; Holtz et al. 2004; Smith et al. 2007; Benson, 2010a), and Poekilopleuron has been considered a spinosauroid (Allain 2002; Allain & Chure 2002) and an allosauroid (Benson 2010a; Benson et al. 2010). Many other taxa are quite fragmentary, and have either been excluded from most previous analyses or remained ambiguously resolved (e.g. Duriavenator, Magnosaurus, Marshosaurus) (Fig. 5). In summary, although a great deal of progress has been achieved in recent years (measured mainly by increased consensus), several points of uncertainty remain in tetanuran phylogeny and are therefore of primary interest here. These are: (1) whether spinosauroids (= megalosauroids) and allosauroids form a clade, or are serially arranged

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outside Coelurosauria; (2) whether ‘megalosaurs’ form a valid clade and, if so, its membership; (3) placement of fragmentary forms of potential geographic and temporal importance; and (4) placement of relatively well known but problematical forms (e.g. Cryolophosaurus, Marshosaurus, Monolophosaurus, Neovenator and Piatnitzkysaurus). Resolving these and other phylogenetic problems within Tetanurae will allow a more reliable analysis of the evolutionary history, diversity and biogeography of this important theropod clade.

Problematical taxa Herrerasauridae and Eoraptor The primitive taxon Eoraptor, along with the various purported constituents of the Herrerasauridae (often including Chindesaurus, Herrerasaurus (with junior synonyms Frenguellisaurus and Ischisaurus) and Staurikosaurus; Sereno 1999), have remained controversial since their original descriptions. Despite the availability of complete materials of both Eoraptor and Herrerasaurus, significant disagreement remains regarding their phylogenetic placement. These forms have been hypothesized as basal theropods (Sereno et al. 1993; Sereno 1999, 2007; Ezcurra & Novas 2007; Nesbitt et al. 2009), basal saurischians (Langer 2004; Irmis et al. 2007), basal dinosaurs and dinosauriformes (Fraser et al. 2002). This paper does not address this question directly, instead focusing on the relationships of definitive theropod taxa. We merely assume that both Eoraptor and Herrerasaurus are more primitive than all the ingroup taxa included in this analysis, and thus can be used to polarize character states for these forms. Whether Eoraptor and Herrerasaurus lie within or outside Theropoda is not strictly relevant to their utility in this regard.

Coelophysoids and ceratosaurs The taxonomic and systematic history of many of these forms has been reviewed in detail elsewhere (Rauhut & Hungerbühler 2000; Carrano & Sampson 2004, 2008; Tykoski & Rowe 2004) and will only be summarized here. Coelophysoids are now understood to represent a basal radiation of theropods, globally distributed but confined to the Late Triassic and Early Jurassic (Carrano & Sampson 2004; Tykoski & Rowe 2004). Once linked as a sister group to ceratosaurs (Rowe & Gauthier 1990; Sereno 1999; Tykoski & Rowe 2004), most workers now support a more basal placement for coelophysoids (Forster 1999; Sampson et al. 2001; Carrano et al. 2002; Rauhut 2003; Smith et al. 2007; Benson 2010a). Correspondingly, true ceratosaurs are more closely related to tetanurans than to coelophysoids. Their record extends from the Early Jurassic through the Late Cretaceous (Carrano & Sampson 2008).

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Recent work has called the monophyly of Coelophysoidea into question (Rauhut 2003; Smith et al. 2007; Nesbitt et al. 2009), but these results are controversial and do not bear directly on the issues addressed in our study.


Megalosaurs and spinosaurs Megalosaurus and its taxonomic descendants have been the most persistently problematic theropod group almost since the naming of the genus in 1824 (Buckland 1824). Like other early named dinosaur genera (e.g. Iguanodon, Plateosaurus), the genus Megalosaurus came to house a wide array of species (in this case, 48) spanning nearly the entire Mesozoic and almost every continent. Unfortunately, the type species M. bucklandii was based on a series of disarticulated and largely disassociated bones from several quarries around the village of Stonesfield, in the recurrent ‘Stonesfield Slate’ facies of the Taynton Limestone Formation (Bathonian, Middle Jurassic; Boneham & Wyatt 1993). Until recently (Benson et al. 2008; Benson 2009a, b, 2010a), this taxon had not been adequately diagnosed, nor had its hypodigm been thoroughly delineated or justified. As a result, it was impossible to clarify what clade, if any, might be based on Megalosaurus. This problem had attendant nomenclatural difficulties, because several higher level taxa (Megalosauridae, Megalosauroidea) are founded on the genus Megalosaurus and would have priority over more recent terms should that genus prove valid. Spinosaurus and similar forms have likewise posed taxonomic and phylogenetic problems since their initial discovery (Stromer 1915; Charig & Milner 1986). The unusual nature of spinosaurid vertebrae and jaws led to widely diverging opinions about their relationships. Spinosaurus was initially given its own family (Stromer 1915; Huene 1926b), within which it was sometimes linked to other high-spined taxa (Acrocanthosaurus, Altispinax) (Walker 1964; Romer 1966; Steel 1970; Carroll 1988), but it has also been placed among the megalosaurids (Romer 1956; Swinton 1970). The more recent discovery of Baryonyx along with newer specimens of Spinosaurus led to debate concerning the potential relationships of these two forms (Charig & Milner 1986, 1990, 1997; Paul 1988a; Buffetaut 1989, 1992; Buffetaut & Ouaja 2002). They are now considered to represent subfamilies within a monophyletic Spinosauridae (Sereno et al. 1994, 1998) that excludes both Acrocanthosaurus and Altispinax. Furthermore, recent discoveries (Galton & Jensen 1979; Allain 2002) and phylogenetic work (Bakker et al. 1992; Sereno et al. 1994, 1998; Holtz 2000; Allain 2002; Benson 2010a) have indicated that ‘megalosaurs’ and spinosaurids probably form a larger clade. It remains to be determined whether other more fragmentary forms can be placed within this group, which has been previously defined on a relatively small number of synapomorphies (Sereno et al. 1994, 1998).

Materials and methods Outgroup relationships Eoraptor and Herrerasaurus serve as outgroup taxa. In choosing multiple outgroups, we allow for greater resolution of character states at root nodes (Barriel & Tassy 1998). These two taxa have been considered basal to all other theropods in all recent analyses, regardless of whether they have been placed as true theropods (Sereno et al. 1993; Sereno 2007; Nesbitt et al. 2009), basal saurischians (Langer 2004) or outside Dinosauria (Fraser et al. 2002).

Operational taxonomic units This study focuses on the relationships of basal tetanurans, and so most of the operational taxonomic units (OTUs) have at least occasionally been referred to this group. This includes forms often referred to as megalosaurs, spinosaurs and allosaurs, and represents the majority of basal tetanuran taxa. Several taxa whose placement had not previously been addressed, or which remained uncertain, were also included (e.g. Cryolophosaurus, Leshansaurus, Marshosaurus, Piatnitzkysaurus, Shidaisaurus). In order to explore whether any of these forms had closer affinities with other clades of basal theropods, we included multiple OTUs from two major non-tetanuran clades. The first, Coelophysoidea, is typically considered the most basal neotheropod clade and is here documented by Dilophosaurus and two species of Coelophysis; its monophyly has recently been questioned (Nesbitt et al. 2009). The second, Ceratosauria, has recently been hypothesized to occupy a more derived position than coelophysoids (e.g. Carrano & Sampson 2008), and is represented here by Elaphrosaurus, Ceratosaurus, Masiakasaurus and Majungasaurus. These taxa were chosen both to represent ingroup character states for these clades (without resorting to the use of suprageneric taxa), and to reflect known state variations within each group. Ornitholestes, Proceratosaurus and Compsognathus were selected to represent relatively completely known but less derived members of Coelurosauria (e.g. Holtz 2000). That is, these taxa are not considered to be derived ornithomimosaurs, oviraptorosaurs, deinonychosaurs etc., and therefore exhibit at least some primitive coelurosaurian character states. In total, the ingroup consisted of 59 OTUs. Our goal in selecting these taxa was to effect a compromise between maximal inclusiveness and productive analysis. Numerous basal tetanuran taxa are so fragmentary and preserve so few codable characters that their inclusion would swamp the matrix with missing data. However, we also deem it insufficient to exclude taxa simply on the basis that they are not complete. Such forms may preserve important data, including unique combinations of character states, which significantly impact the phylogenetic results. We



also include several taxa that cannot be diagnosed based on unique character states, but which we consider are likely to represent valid species given their temporal and/or geographic situation. Ingroup taxa are listed below, including information on hypodigm, provenance, and diagnostic features. The diagnoses presented below combine original and (where cited) previously published observations. They are contextualized within that clade for which most of the listed features are unique. For currently non-diagnosable taxa (sensu stricto), we include comments on the potential viability of the form in lieu of a traditional diagnosis. Specimens in bold were examined by the authors firsthand; the remainder were studied via published photographs and drawings.

Institutional abbreviations AM: Albany Museum, Grahamstown; AMNH: American Museum of Natural History, New York; AODF: Australian Age of Dinosaurs Museum of Natural History, Winton; BP: Bernard Price Institute, Johannesburg; BSP: Bayerische Staatssammlung für Paläontologie und Historische Geologie, München; BYU: Brigham Young University, Provo; CEPUNL: Centro de Estratigrafia e Paleobiologia da Universidade Nova de Lisboa, Lisboa; CM: Carnegie Museum of Natural History, Pittsburgh; CMP: Cantera Mas de la Parreta collection, Museo de Valltorta, Tirig; CPP: Centro de Pesquisas Paleontológicas Llewellyn Ivor Price, Peirópolis; CV: Municipal Museum of Chongqing, Chongqing; DINO/DNM: Dinosaur National Monument, Vernal; DMNH: Denver Museum of Nature and Science, Denver; DORCM: Dorset County Museum, Dorchester; DZSWS: Devizes Museum, Devizes; FMNH: Field Museum of Natural History, Chicago; FPDM: Fukui Prefectural Dinosaur Museum, Katsuyama; FSL: Faculté des Sciences, Université Claude Bernard, Lyon; GA: Sección de Geolog´ıa, Sociedad de Ciencias Aranzadi, San Sebastián; GC: Goucher College, Baltimore; HASMG: Hastings Museum and Art Gallery, Hastings; IFPUB: Institut für Geologische Wissenschaften der FU Berlin, Fachbereich Paläontologie, Berlin; IMGP: Institut und Museum für Geologie und Paläontologie, Georg-August-Universität, Göttingen; IPS/IPSN: Institut Paleontològic Dr. Miquel Crusafont, Sabadell; IWCMS: Museum of Isle of Wight Geology, Sandown; IVPP: Institute of Palaeontology and Palaeoanthropology, Beijing; KMV: Kunming Municipal Museum, Kunming; KPE: Earth Science Department, Kyungpook National University, Daegu; KS: Kyeongnam Science High School Museum, Kyeongnam; LDMLCA: Lufeng Dinosaur Museum-Lufeng Chuanjie A’na, A’na; MACN: Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires; MB: Humboldt Museum für Naturkunde, Berlin; MCCM: Museo de las Ciencias de Castilla-La Mancha, Cuenca; MCF: Museo Municipal ‘Carmen Fuñes’, Plaza Huincul; MCZ: Museum of Comparative Zoology, Harvard University, Cambridge;

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MG/MGSP: Museu Geológico, Lisbon; MH: Naturhistorisches Museum, Basel; MIWG: ‘Dinosaur Isle’ Museum of Isle of Wight Geology, Sandown; ML: Museu da Lourinhã, Lourinhã; MLP: Museo de La Plata, La Plata; MM: Université de Montpellier, Montpellier; MMR/UFU: Museu de Minerais e Rochas ‘Heinz Ebert,’ Rio Claro; MN: Museu Nacional/Universidad Federal do Rio de Janeiro, Rio de Janeiro; MNHN: Muséum National d’Histoire Naturelle, Paris; MNHNUL: Museu Nacional de História Natural da Universidade de Lisboa, Lisbon; MNN: Musée National du Niger, Niamey; MOAL: Molina Alto collection, Museo de la Fundación Conjunto Paleontológico de Teruel-Dinópolis, Teruel; MOR: Museum of the Rockies, Bozeman; MPCA: Museo Provincial ‘Carlos Ameghino’, Cipoletti; MPEF: Museo Paleontológico ‘Egidio Feruglio’, Trelew; MPM: Museo Padre Molina Paleontolog´ıa de Vertebrados, R´ıo Gallegos; MPZ: Museo Paleontológico de la Universidad de Zaragoza, Zaragoza; MSNM: Museo delle Scienze Naturale di Milano, Milano; MUCP: Museo de la Universidad Nacional del Comahue, Neuquén; MWC: Museum of Western Colorado, Fruita; NCSM: North Carolina Museum of Natural Sciences, Raleigh; NHMUK: Natural History Museum, London (formerly BMNH); NMV: National Museum of Victoria, Melbourne; OMNH: Oklahoma Museum of Natural History, Norman; OUMNH: Oxford University Museum, Oxford; PIN: Paleontological Institute, Russian Academy of Sciences, Moscow; PS: Colectivo Arqueológico-Paleontológico de Salas, Salas de los Infantes; PVL: Fundación Miguel Lillo, Universidad Nacional de Tucumán, San Miguel de Tucumán; PW: Paleontological Collections, Department of Mineral Resources, Bangkok; QG: National Museum of Natural History, Bulawayo; QW: Giant Buddha Temple Museum, Leshan; ROM: Royal Ontario Museum, Toronto; SDM: Stroud and District Museum, Dorset; SDSM: South Dakota ´ School of Mines, Rapid City; SGM: Ministère de l’Energie et des Mines, Rabat; SM: Sirindhorn Museum of Palaeontology, Sahat Sakhan; SM∗ : Senda Miravete collection, Museo de la Fundación Conjunto Paleontológico de TeruelDinópolis, Teruel; SMA: Sauriermuseum Aathal, Aathal; SMNS: Staatliches Museum für Naturkunde, Stuttgart; SMU: Southern Methodist University, Dallas; SNGM: Servicio Nacional de Geológia y Minero, Santiago de Chile; TATE: Tate Museum, Casper College, Casper; TUB: Technishe Universität Berlin, Berlin; UA: Département de Paléontologie, Université d’Antananarivo, Antananarivo; UCMP: University of California Museum of Paleontology, Berkeley; UC OBA: Department of Organismal Biology and Anatomy, University of Chicago, Chicago; UCPC: University of Chicago Paleontological Collection, Chicago; UMNH/UUVP: Natural History Museum of Utah (formerly Utah Museum of Natural History) University of Utah, Salt Lake City; UOP: University of Portsmouth, Portsmouth; USNM: National Museum of Natural History, Smithsonian Institution, Washington; USP: Universidade

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de São Paulo, São Paulo; UT: Geology Department, University Al Fateh, Tripoli; WAM: Western Australia Museum, Perth; YNUGI: Geological Institute, Yokohama National University, Yokohama; YPM: Peabody Museum of Natural History, Yale University, New Haven; ZDM: Zigong Dinosaur Museum, Zigong; ZIN: Zoological Institute, Russian Academy of Sciences, St. Petersburg.

Systematic palaeontology Acrocanthosaurus atokensis Stovall & Langston, 1950


1950 Acrocanthosaurus atokensis Stovall & Langston: 686, figs 2–4, pls 1–3. Holotype. OMNH 10146 (= MUO 8-0-S9), an incomplete skull and partial postcranial skeleton (Stovall & Langston 1950). Hypodigm. Holotype; OMNH 10147 (= MUO 8-0-S8), a partial skeleton (Stovall & Langston 1950); NCSM 14345, a nearly complete skull and skeleton (Currie & Carpenter 2000); and SMU 74646, an incomplete skull and skeleton (Harris 1998a). Diagnosis. Allosauroid theropod with: (1) absence of nasal extension of antorbital fossa and associated pneumatopores; (2) supraoccipital expanded parasagittally into double boss posterior to nuchal crest (Currie & Carpenter 2000); (3) cervical vertebral neural spines with triangular anterior processes that insert into fossae ventral to overhanging processes on preceding neural spines (Harris 1998a); (4) neural spines of presacral, sacral and anterior caudal vertebrae more than 2.5 times taller than respective centrum lengths (Stovall & Langston 1950); and (5) accessory process on lateral surface of caudal prezygapophysis (Stovall & Langston 1950). Occurrence. McLeod Prison, Arnold Farm and Cochran Farm, Atoka and McCurtain Counties, Oklahoma, and Hobson Ranch, Parker County, Texas, USA; Antlers and Twin Mountains Formations; late Aptian–early Albian, Early Cretaceous. Remarks. Discovered more than half a century ago, Acrocanthosaurus has occupied many systematic positions among the large-bodied ‘carnosaurs’ since then, with various workers suggesting affinities with allosaurids (Stovall & Langston 1950; Colbert & Russell 1969), spinosaurids (Walker 1964; Romer 1966; Steel 1970; Carroll 1988) and megalosaurids (Romer 1956; Swinton 1970; Russell 1984). More recently, consensus has favoured placement of Acrocanthosaurus within Allosauroidea, and debate has focused on whether its closest ingroup affinities lie with Carcharodontosauridae (Sereno et al. 1996; Harris 1998a; Sereno 1999; Brusatte & Sereno 2008; Benson 2010a; Benson et al.

2010; Eddy & Clarke 2011) or Allosauridae (Holtz 1994a, 2000; Currie & Carpenter 2000; Novas et al. 2005; Smith et al. 2007). Notably, the eponymous ‘high spined’ dorsal vertebrae have deep lateral embayments, resulting in an ‘I-beam’ cross-section that is shared with Giganotosaurus and Mapusaurus (Coria & Currie 2006; Benson 2010a). Two recently described specimens (Harris 1998a; Currie & Carpenter 2000; Coria & Currie 2006; Eddy & Clarke 2011) essentially complete the known osteology of this animal, allowing a thorough assessment of its systematic morphology. Like Giganotosaurus and Carcharodontosaurus, these recent finds have demonstrated the gigantic size of this taxon, which approached the dimensions (although not the mass, based on femoral circumference) of the largest tyrannosaurids. Aerosteon riocoloradensis Sereno et al., 2008 2008 Aerosteon riocoloradensis Sereno et al.: 4, figs 2–11, 12A, 13–16. Holotype. MCNA-PV-3137, an incomplete skull and skeleton. Diagnosis. Allosauroid theropod with: (1) robust, cylindrical transverse processes on proximal caudal vertebrae; and (2) fossa on lateral surface of coracoid dorsal to glenoid and (separate) subglenoid fossa. Occurrence. Cañadon Amarillo, Mendoza, Argentina; Anacleto Formation, R´ıo Colorado Subgroup, Neuquén Group; early Campanian, Late Cretaceous (Dingus et al. 2000; Leanza et al. 2004). Remarks. Sereno et al. (2008) were uncertain of the affinities of Aerosteon within Tetanurae. Consequently, they listed several features in its diagnosis that can now be identified in Neovenator (pneumatic canal within posterior dorsal transverse processes), Acrocanthosaurus (dorsal neural spines with central pneumatic space), Orkoraptor (short ventral process of prefrontal) and other allosauroids (anterior dorsal vertebra with very large parapophyses; anterodorsally inclined posteriormost dorsal neural spine; pneumaticity in caudal vertebral centra), or simply cannot be compared in closely related taxa. Based on recent comparative work (Benson et al. 2010), our emended diagnosis comprises two probable autapomorphies that are absent in all other allosauroids, including a likely sister taxon of Aerosteon, Megaraptor. Afrovenator abakensis Sereno et al., 1994 1994 Afrovenator abakensis Sereno et al.: 270, fig. 2. Holotype. MNN TIG1 (= UC OBA 1), an incomplete skull and skeleton.

The phylogeny of Tetanurae Diagnosis. Megalosauroid theropod with: (1) rounded, lobate anterior maxillary margin of antorbital fossa (Sereno et al. 1994); (2) low, rectangular neural spine on third cervical vertebra (Sereno et al. 1994); (3) broad lateral flange on metacarpal I for articulation with metacarpal II (Sereno et al. 1994); and (4) posterior extension of distal pubic expansion inset from lateral surfaces of anterior extension, forming pair of conjoined, posterior flanges.


Occurrence. In Abaka, Agadez, Niger; Tiourarén Formation, Irhazer Group; Middle–Late Jurassic or Neocomian, Early Cretaceous (Lapparent 1960; Sereno et al. 2004; Rauhut & López-Arbarello 2009). Remarks. Afrovenator is represented by one of the most complete theropod specimens known from Africa, including portions of all major skeletal regions. Originally described as a basal torvosauroid (megalosauroid) (Sereno et al. 1994), it has also been recovered as an allosauroid (Rauhut 2003). Afrovenator exhibits a number of features that seem to ally it with megalosaurs or other basal tetanurans. In particular, the long, low skull shows little ornamentation, and the pelvic girdle more closely resembles that of Marshosaurus and Piatnitzkysaurus than those of allosauroids in the relative sizes of the iliac peduncles and morphology of the distal pubis. The maxilla bears a long anterior ramus, a ‘kinked’ ascending ramus and a large imperforate maxillary ‘fenestra’ resembling those of megalosaurs such as Marshosaurus (DNM 343/DINO 16455b), Duriavenator and Eustreptospondylus. Genus Allosaurus Marsh, 1877 1877 Allosaurus Marsh: 515. 1878 Creosaurus Marsh: 243, fig. 1. 1878 Epanterias Cope: 406. 1879 Labrosaurus Marsh: 91. Included species. Allosaurus europaeus Mateus et al., 2006; Allosaurus fragilis Marsh, 1877 and Allosaurus jimmadseni Chure et al., 2006 (nomen nudum). Hypodigm. A. europaeus: ML 415 (holotype). A. fragilis: YPM 1930 (holotype) and AMNH 275, 287, 290, 324, 408, 496, 600, 666, 680, 813, 851, 5750, 5753, 5767 (holotype, Epanterias amplexus), 6125, 6128; BYU 2028, 4861, 5164, 5268, 5292, 571-8901, Mes 5583, 11936, 13621, 16942, 17106, 17281; CM 11844; DMNH 44397; DINO 3984; FMNH 1505, P25114; MCZ 3897; ROM 12868; UMNH VP 1251, 3113, 5316, 5326–5328, 5470, 5480, 6317, 6340, 6365, 6400, 6408, 6473, 6475, 6499, 6502, 7190, 7408, 7411, 7794, 7880, 7882, 7884–7885, 7889–7891, 7895, 7898, 7908, 7922, 7926–7930, 7932, 7934, 7937–7938, 7957, 7966, 8102, 8123, 8142, 8151, 8229, 8240–8241, 8355, 8397, 8484, 9103, 9147, 9149, 9162, 9168, 9180, 9191, 9201, 9212, 9323, 9327, 9366, 9376, 9401, 9470,

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9473, 9480, 9500, 9502, 9505, 9514, 9709, 10360, 10386, 10779, 11031, 11463, 12231, 16584–16585; UUVP 1403, 3304, 3894, 3981, 6000 (‘neotype’ = DINO 2560); USNM 2323, 3304, 4734 (topotype, probably also including USNM 2315, holotype, Labrosaurus ferox), 4737, 7336, 8257, 8302, 8335, 8367, 8405, 8423; YPM 1879?, 1890 (holotype, Creosaurus atrox), 1931 (holotype, Labrosaurus lucaris). A. jimmadseni: DINO 11541; MOR 693; SDSM 30510; SMA 0005; UMNH VP C481. Diagnosis. Allosauroid theropod with: (1) tall, mediolaterally compressed dorsal projection (‘horn’) on posterodorsal surface of lacrimal (Chure 2001a); (2) reduced external mandibular fenestra; (3) strongly downturned paraoccipital processes that terminate well ventral to basal tubera; (4) neomorphic antarticular bone in lower jaw (Madsen 1976a); and (5) distal expansion of ischium suboval in lateral view (modified from Madsen 1976a). Occurrence. Allosaurus jimmadseni and A. fragilis are known from dozens of sites in Utah, Arizona, New Mexico, Colorado, Wyoming, Montana and South Dakota, USA (cf. Turner & Peterson 1999; Foster 2003); Salt Wash and Brushy Basin Members, Morrison Formation; Kimmeridgian–Tithonian, Late Jurassic. A. europaeus derives from Andrés, Praia de Vale Frades and Guimarota, Leiria, Portugal; Alcobaça and Porto Novo Members, Lourinhã Formation; late Kimmeridgian–Tithonian, Late Jurassic (Pérez-Moreno et al. 1999; Rauhut & Fechner 2005; Mateus et al. 2006). Remarks. One of the most abundant large theropods in the world, and consequently one of the best known, Allosaurus has been the subject of two monographic descriptions (Gilmore 1920; Madsen 1976a) and numerous studies on its growth and ontogeny (Bybee et al. 2006; Loewen 2009), population variation (Chure & Madsen 1996; Smith 1998), pathologies (Chure 2000; Hanna 2002) and taxonomy (Smith 1998; Chure 2001a). In the last century it has also largely occluded Megalosaurus as an archetypal ‘basic’ theropod. Most of this material pertains to A. fragilis, although the recent discovery of a second North American species has necessitated the transfer of some specimens into the new taxon. A. jimmadseni is based on a nearly complete skull and skeleton from Dinosaur National Monument (Chure 2000). It was mentioned in print (Chure et al. 2006) but has been described in detail only in a dissertation (Chure 2001a) and so is technically a nomen nudum. A. jimmadseni is approximately equivalent to the ‘creosaur-type allosaurid’ described by Bakker (2000; which does not include the type of Creosaurus). Because descriptive research on A. jimmadseni is ongoing (D. Chure pers. comm.; M. Loewen pers. comm.), we do not discuss its morphology in detail but include it in the hypodigm of Allosaurus.

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A. europaeus was described on the basis of a partial skull (ML 415) by Mateus et al. (2006) that has also been regarded as belonging to A. fragilis (Malafaia et al. 2007); certainly it is more similar to that species than to A. jimmadseni. Other Allosaurus material from Portugal (MNHNUL/AND.001, a partial pelvis and hind limb) was referred to A. fragilis (Pérez-Moreno et al. 1999) and A. europaeus (Mateus et al. 2006) but may not be determinate to the species level and is here regarded as Allosaurus sp. A juvenile maxilla from Guimarota (IFPUB Gui Th 4) has also been assigned to Allosaurus sp. (Rauhut & Fechner 2005). Regardless of the validity of A. europaeus, we consider these specimens to be correctly assigned to the genus Allosaurus. Angaturama limai Kellner & Campos, 1996 1996 Angaturama limai Kellner & Campos: 152, figs 2–4. Holotype. USP GP/2T-5, the anteriormost portion of a rostrum. Diagnosis. Spinosaurid with midline ridge on dorsal surface of conjoined premaxillae extending further anteriorly than in Baryonyx and Suchomimus (modified from Kellner & Campos 1996). Angaturama cannot currently be distinguished from Irritator because it is based on nonoverlapping skull material. Occurrence. Unspecified locality in the Araripe Basin, southern Ceará, Brazil; Romualdo Member, Santana Formation; Albian, Early Cretaceous (Kellner & Campos 1996). Remarks. Angaturama possesses Spinosaurus-like dental characters such as the absence of serrations on the carinae. It also shows the presence of a midline crest on the dorsal surface of the premaxilla, similar to those of Baryonyx and Suchomimus. However, it differs from other spinosaurids in that no other taxon shows both features in combination. Angaturama is based on very fragmentary remains that may represent the same taxon, or perhaps the same individual, as the holotype of Irritator (Sereno et al. 1998; Sues et al. 2002; Dal Sasso et al. 2005). However, this remains speculative until the two holotypic specimens can be compared directly, and we retain the two taxa as distinct OTUs. Australovenator wintonensis Hocknull et al., 2009 2009 Australovenator wintonensis Hocknull et al.: 24, figs 20–37. Holotype. AODF 604, a partial skeleton lacking most of the skull.

Diagnosis. Allosauroid with: (1) lateral groove along ulnar shaft (Hocknull et al. 2009); and (2) anterior bevelling of proximal fibular articular surface (Hocknull et al. 2009) Occurrence. AODF 85, ‘Matilda Site,’ Elderslie Station, 60 km north-west of Winton, Queensland, Australia; Winton Formation; late Albian, Early Cretaceous (Hocknull et al. 2009). Remarks. The recently described holotype of Australovenator is the most complete theropod skeleton known from Australia. The original description was accompanied by a phylogenetic analysis that placed it within Allosauroidea, close to Neovenator but just outside Carcharodontosauridae. Indeed, although the skull is incomplete, the left dentary is quite similar to that of Neovenator in its slender proportions and the arrangement of neurovascular foramina; these taxa also share extensively pneumatized ribs (among allosauroids). A number of other features listed in the original diagnosis of Australovenator can be found in other theropods, particularly Megaraptor, Chilantaisaurus and Fukuiraptor, suggesting that all these taxa may share a close phylogenetic relationship (Benson et al. 2010). Baryonyx walkeri Charig & Milner, 1986 1986 Baryonyx walkeri Charig & Milner: 359, figs 1–4. Holotype. NHMUK R9951, a partial skull and skeleton. Hypodigm. Holotype and UOP C001.2004, cast of a dorsal vertebra. Diagnosis. Spinosaurid theropod with: (1) midline knob at posterior end of conjoined nasals terminating in cruciate process (Charig & Milner 1986); (2) subrectangular lacrimal horn (Sereno et al. 1998); and (3) peg-and-notch articulation between scapula and coracoid (Sereno et al. 1998). Occurrence. Smokejacks Brickworks (Ockley brick pit), Walliswood, Ockley, near Dorking and Ewhurst Brickworks, Surrey, and south-west coast of the Isle of Wight, England; Cypridea clavata zone, Upper Weald Clay and Wessex Formations; Barremian–early Aptian, Early Cretaceous (Charig & Milner 1997; Hutt & Newberry 2004). Remarks. The discovery of Baryonyx represented the first reasonably complete theropod from the English Wealden in more than a century and a half of collecting. It also became the best known spinosaurid until the discovery of Suchomimus (Sereno et al. 1998). The unusual and highly derived nature of the skull and skeleton led to initial controversies surrounding its anatomy and relationships (Charig

The phylogeny of Tetanurae & Milner 1986, 1990, 1997; Paul 1988a; Sereno et al. 1998), but subsequent finds have supported its placement within the Spinosauridae (sometimes within its own subfamily, Baryonychinae; Sereno et al. 1998; Holtz et al. 2004; Benson 2010a) and have clarified the identities and details of several skull bones. Genus Carcharodontosaurus Stromer, 1931 1931 Carcharodontosaurus Stromer: 19, pl. 1, figs 1–15.


Included species. C. saharicus (Depéret & Savornin, 1925) Stromer, 1931 and C. iguidensis Brusatte & Sereno, 2007. Hypodigm. C. saharicus: BSP 1922 X46 (‘genoholotype’; destroyed), a partial skull and skeleton, 1922 X45 (‘Spinosaurus B’; destroyed), cervical vertebrae and pedal phalanges; MB.R.2056, endocast from BSP 1922 X46; IMGP 969-1/2/3; SGM Din-1, partial skull (neotype), UCPC OT6, cervical vertebra, Din-3, cervical vertebra. C. iguidensis: MNN IGU2, a left maxilla (holotype) and IGU3–6, 11, teeth, skull elements, and a cervical centrum. Diagnosis. Allosauroid theropod with: (1) large paracondylar and internal carotid artery pneumatic recesses (Brusatte & Sereno 2007); (2) funnel-shaped basisphenoid recess (Brusatte & Sereno 2007); (3) reniform posterior centrum face in postaxial cervicals, twice as wide as tall (Sereno et al. 1996); and (4) deep ventral keel on postaxial cervicals that approaches depth of centrum (Sereno et al. 1996). Occurrence. C. saharicus is known from Timimoun, Algeria, the Baharˆıje Oasis, Egypt and the Kem Kem region, Morocco; ‘Continental Intercalaire,’ Baharˆıje, and Kem Kem beds, respectively; Albian?–Cenomanian, late Earlyearly Late Cretaceous (Stromer 1931; Lapparent 1960; Brusatte & Sereno 2007). C. iguidensis is known from Iguidi, west of In Abangarit, Agadez, Niger; Echkar Formation, Tegama Group; Cenomanian, early Late Cretaceous (Brusatte & Sereno 2007). Remarks. Like many large Jurassic and Cretaceous theropods, Carcharodontosaurus was originally described as a species of Megalosaurus (M. saharicus, Depéret & Savornin 1925) and has the rather unusual distinction of having been referred to two different species of this taxon (M. africanus, Huene 1956, p. 489 [incorrectly attributed by that author to Depéret & Savornin 1925 and possibly a lapsus calami]). In a later paper (Depéret & Savornin 1928), the original authors suggested an affiliation with the North American genus Dryptosaurus, but since Rauhut (1995) Carcharodontosaurus has uniformly been recognized as an allosauroid (e.g. Sereno et al.

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1996; Brusatte & Sereno 2007). The syntype series of M. saharicus consists of two teeth. However, despite his reservations about the diagnostic utility of isolated theropod teeth, Stromer (1931, p. 10) specified this taxon as the type species of the new genus Carcharodontosaurus, designating a more complete specimen from Egypt as the ‘genoholotype’ (Stromer 1931; reviewed by Rauhut 1995). The syntype teeth are now considered lost and Brusatte & Sereno (2007) designated a partial skull (SGM Din1) from the upper Kem Kem beds, penecontemporaneous with the Baharˆıje Beds in Egypt, as the neotype of C. saharicus; it was claimed that “these Egyptian fossils were never cast” (Brusatte & Sereno 2007, p. 904) but in fact an endocast from BSP 1922 X46 was made and currently resides in Berlin as MB.R.2056. The neotype includes much of the skull and indicates an individual of giant size. Using a reconstructed premaxilla based on those of Giganotosaurus and Mapusaurus, we estimate the skull length of this specimen as close to 142 cm, equivalent to large specimens of Tyrannosaurus rex (e.g. FMNH PR 2081, at 139 cm). The femoral cross sectional proportions of BSP 1922 X46 suggests an animal of lower total mass than a Tyrannosaurus of approximately equal femoral length. Largely because of the distinctive crenulations present on the teeth of Carcharodontosaurus, the genus has been identified across mid-Cretaceous deposits of North Africa, primarily from the ‘Continental Intercalaire’ and its descendant units; these beds vary in age from Albian through Cenomanian. The characteristic crenulations of isolated ‘Carcharodontosaurus sp.’ teeth are present in other derived carcharodontosaurids such as Giganotosaurus (MUCPvCh 1) and Mapusaurus (Coria & Currie 2006), vary in prominence between individuals and along the tooth row, and are observed in a lesser developed form in numerous other tetanuran taxa (Brusatte et al. 2007). Given the recent identification of a second species from Niger (Brusatte & Sereno 2007), we consider it prudent to refer such isolated teeth to cf. Carcharodontosauridae indet. In addition, a complicated debate has surrounded the identity of two fragmentarily known forms from North Africa. The first, called ‘Spinosaurus B’ by Stromer (1934, p. 21), is represented by a vertebral series from the Baharˆıje Beds of Egypt. The second specimen, based on several isolated cervical vertebrae from the Kem Kem Beds of Morocco, was described as Sigilmassasaurus brevicollis (Russell 1996). Sereno et al. (1996) referred both to C. saharicus. However, this was disputed by Rauhut (2003) and Novas et al. (2005), who claimed that although the vertebrae of ‘Spinosaurus B’ and Sigilmassasaurus resembled one another, they were distinct from those associated with the Carcharodontosaurus vertebrae from BSP 1922

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X46. The issue warrants more extended treatment elsewhere, and at present we do not use specimens of Sigilmassasaurus for our character codings.

and metatarsals; these likely pertain to the same individual as the lectotype (Hu 1964; Benson & Xu 2008).

Ceratosaurus nasicornis Marsh, 1884

Diagnosis. Tetanuran theropod with: (1) subrectangular, anteromedially curving humeral deltopectoral crest protruding almost as far anteriorly as it is long proximodistally, and bearing pitted scar on anterior surface (Rauhut 2003; Benson & Xu 2008); and (2) obliquely oriented radial (lateral) condyle of humerus (Benson & Xu 2008).

1884 Ceratosaurus nasicornis Marsh: 330, pls 8–14. 1892 Megalosaurus nasicornis Cope: 241. 2000 Ceratosaurus magnicornis Madsen & Welles: 2, figs 1, 3, pls 1–8. 2000 Ceratosaurus dentisulcatus Madsen & Welles: 21, figs 2–5, 10, pls 9–23


Holotype. USNM 4735 (= YPM 1933), a complete skull and partial skeleton. Hypodigm. Holotype and MWC 1.1 (holotype, Ceratosaurus magnicornis), complete skull and partial skeleton; UMNH VP 5278 (holotype, Ceratosaurus dentisulcatus), partial skull and skeleton; BYU-VP 5010, left metatarsal III; 5008, left metatarsal III; 4838, 4853, 4908, 4968, 5092, 5132, 5133, 5135, 8937, 8938, 8974, 8982, 9099, 9108, 9141, 9152, 9161–9163, 9165, caudal vertebrae; 4951, 4952, 8907, 9142–9144, dorsal vertebrae; 12893, skull, vertebrae and partial femur; 17550, articulated pelvis and sacrum. Diagnosis. Ceratosaur with: (1) mediolaterally narrow, rounded midline horn core on nasals, fused in adults (modified from Marsh 1884); (2) medial oval groove on nasals behind horn core (Rauhut 2003); (3) pubis with large, rounded notch ventral to obturator foramen (Rauhut 2003); and (4) small median dorsal osteoderms (Marsh 1884). Occurrence. Marsh-Felch Quarry, Cañon City, Fremont County, Fruita area, Mesa County and Dry Mesa Quarry, Montrose County, Colorado; Cleveland-Lloyd Quarry and San Rafael Swell, Emery County, and Dinosaur National Monument, Uintah County, Utah; Quarry 9, Como Bluff, Albany County, Wyoming, USA; lower Brushy Basin Member, Morrison Formation; Kimmeridgian–Tithonian, Late Jurassic (Turner & Peterson 1999, 2004; Madsen & Welles 2000). Remarks. This taxon has been reviewed recently (Carrano & Sampson 2008), and we only reiterate that we consider the existing North American specimens of Ceratosaurus to represent a single species, C. nasicornis. Chilantaisaurus tashuikouensis Hu, 1964 1964 Chilantaisaurus tashuikouensis Hu: 56, figs 1–8. Lectotype. IVPP V.2884.1, a right humerus. Hypodigm. Lectotype and IVPP V.2884.2–V.2884.7, manual ungual phalanx, left ilium, femora, tibiae, left fibula,

Occurrence. Tashuikou (Dashuigou), 60 km north of Chilantai (Jilantai), eastern Alashan Desert, Nei Mongol Zizhiqu, China; Ulansuhai Formation; Turonian or younger (< 92 Ma), Late Cretaceous (Kobayashi & Lü 2003). Remarks. This taxon was recently redescribed in detail, revealing several avetheropod synapomorphies (Benson & Xu 2008). Its taxonomic affinities remain uncertain but it is unlikely to belong to a megalosauroid (spinosauroid) as recovered by Rauhut (2003). Instead, Chilantaisaurus shares features with Fukuiraptor (long humerus relative to femur; long deltopectoral crest; tall, narrow proximal dimensions of manual unguals), Aerosteon and Neovenator (preacetabular shelf on ilium) and Australovenator (narrow manual unguals; median ridge on distal tibia) that suggest a close relationship between these taxa (Benson et al. 2010). Chuandongocoelurus primtivus He, 1984 1984 Chuandongocoelurus primtivus He: 40, figs 6–15. Holotype. CCG 20010, a partial postcranial skeleton. Diagnosis. See Remarks. Occurrence. Chuandong, Sichuan, China; Xiashaximiao Formation; Middle Jurassic (He 1984). Remarks. The holotype of Chuandongocoelurus shows a potentially diagnostic character combination as it possesses tetanuran synapomorphies of the ilium (pubic peduncle of ilium considerably larger than ischial peduncle; pubic peduncle approximately 1.34 times as long anteroposteriorly as broad mediolaterally) and tibia (a vertical ‘supraastragalar’ buttress located medially on the anterior surface of the distal end (Rauhut 2003) in combination with a femoral head that is directed approximately 45◦ anteromedially and inclined ventrally, otherwise seen only in nontetanuran theropods. Chuandongocoelurus also possesses a hypertrophied supraacetabular crest, often considered a non-tetanuran feature (e.g. Holtz 2000), but this has also been reported in Monolophosaurus (Zhao et al. 2010). The ilium and some dorsal centra are the only bones preserved in both taxa, and both show the same combination of tetanuran and ‘non-tetanuran’ features. Therefore the two are

The phylogeny of Tetanurae only distinguishable in size: the Chuandongocoelurus holotype represents an individual measuring 36 mm between the pubic and ischial peduncles, whereas this measurement is 105 mm in Monolophosaurus (IVPP 84019). He (1984) referred a second specimen (CCG 20011; vertebrae) to Chuandongocoelurus, but this material represents a substantially larger individual and shows little overlap with CCG 20010 that would allow positive referral to the same taxon. Furthermore, CCG 20011 possesses anteroposteriorly long cervical centra with anteroposteriorly elongate pleurocoelous fossa, similar to those seen in the ceratosaur Elaphrosaurus. It is therefore unlikely that CCG 20011 represents Chuandongocoelurus.


Coelophysis bauri (Cope, 1887a) 1887a Coelurus bauri Cope: 368. 1887a Coelurus longicollis Cope: 368. 1887b Tanystropheus bauri Cope: 226. 1887b Tanystropheus longicollis Cope: 227. 1887b Tanystropheus willistoni Cope: 227. 1889 Coelophysis bauri Cope: 626. 1889 Coelophysis longicollis Cope: 626. 1889 Coelophysis willistoni Cope: 626. 1984 Longosaurus longicollis Welles: 160. 1991 Rioarribasaurus colberti Hunt & Lucas: 195. Neotype. AMNH 7224, nearly complete skull and skeleton. Hypodigm. Neotype; AMNH 2701–2704, 2705–2707 (lectotype, L. longicollis), 2708, 2715, 2717–2720, 2722 (lectotype, C. bauri), 2747, 2753, 7223 (paratype, C. bauri), 7222, 7242, 7243, 7246; CM 31374; DMNH 14729, 22702, 30209, 30596, 32813; MCZ 4326, 4331, 4332; NMMNH P-41416, 41419, 42200, 42351–42354, 42576–42580, 44551–44556, 44802, 46615, 50528–50537, 55336–55346, 55348–55350, 57652, 57653; YPM 5705, 41196, 41197. Occurrence. Ghost Ranch, Abiquiu, Rio Arriba County, New Mexico; Petrified Forest National Park, Arizona; Petrified Forest Member, Chinle Formation; Norian, Late Triassic. Remarks. Although C. bauri appears to be distinct from its close congener, C. rhodesiensis, the many available specimens of this taxon have not been subjected to detailed phylogenetic study for several decades. As a result, it can be differentiated from other coelophysoids but not diagnosed based on autapomorphies. We do not dispute the validity of C. bauri but note that further anatomical study is still needed. The type genus for Coelophysidae and therefore Coelophysoidea, Coelophysis is a well-known small theropod represented by hundreds of individual specimens at the

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Whitaker Quarry belonging to the type species, C. bauri (cf. Rinehart et al. 2009). In recent years, restudy of both C. bauri and Syntarsus rhodesiensis has led to a general agreement that the two taxa are extremely similar and probably represent species of the same genus (Paul 1988a; Downs 2000; Bristowe & Raath 2004). Coelophysis rhodesiensis (Raath, 1969) 1969 Syntarsus rhodesiensis Raath: 1, figs 1–6, pls 1–5. 1988a Coelophysis rhodesiensis Paul: 262. 2001 Megapnosaurus rhodesiensis Ivie et al.: 63. Holotype. QG1, partial postcranial skeleton. Hypodigm. Holotype and QG 3A, 45, 76, 124, 127, 164–165, 169–1103; BP/I/5246, 5278; numerous skulls and skeletons. Diagnosis. Coelophysoid theropod with: (1) blunt, squared anterior margin of antorbital fossa; (2) base of lacrimal vertical ramus width < 30% its height; and (3) maxillary and dentary tooth rows end posteriorly at anterior rim of lacrimal (all from Bristowe & Raath 2004). Occurrence. Maura River, Southcote farm and Spring Grange farm, Nyamandhlovu, Matabeleland North and Chitake River, Mashonaland North, Zimbabwe; area between farms Edelweiss and Welbedacht, Ladybrand District, Free State, South Africa; Forest Sandstone and Upper Elliot Formations; ?Hettangian–Sinemurian, Early Jurassic. Remarks. Coelophysis rhodesiensis is one of the most abundantly preserved theropods, with hundreds of specimens representing dozens of individuals from a range of sizes and growth stages (Raath 1969, 1977; Bristowe & Raath 2004). The material is in an excellent state of preservation, with both articulated and disarticulated individuals, making C. rhodesiensis one of the most completely known early theropod dinosaurs. Compsognathus longipes Wagner, 1861 1861 Compsognathus longipes Wagner: 94, pl. 3. 1972 Compsognathus corallestris Bidar et al.: 2327. Holotype. BSP ASI 563, complete skull and skeleton. Hypodigm. Holotype; MNHN CNJ 79 (holotype, Compsognathus corallestris), complete skull and skeleton; and MB.R.2003.2, cast of holotype counterslab. Diagnosis. Coelurosaur with a possibly unique character combination of: (1) ventral process at posterior end of

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premaxillary body; (2) opisthocoelous cervical vertebrae; (3) metacarpal I less than 1/3 length of metacarpal II; (4) pollex terminates at or below distal end of phalanx I of digit II; and (5) fourth trochanter on femur absent (all from Peyer 2006).

The impression that Becklespinax possesses a combination of tall and short posterior dorsal neural spines (Naish & Martill 2007) similar to those of Concavenator results from a broken anterior neural spine (confirmed by direct observation of NHMUK R1828). Condorraptor currumili Rauhut, 2005a


Occurrence. Reidenburg-Kelheim area, Bayern, Germany and near Bessons, Petit Plan de Canjuers, Var, ProvenceAlpes-Côte d’Azur, France; Ober Solnhofen Plattenkalk and Canjuers Lithographic Limestone; Tithonian, Late Jurassic (Bidar et al. 1972). Remarks. Two nearly complete specimens document most of the osteology of Compsognathus, and both have been re-examined in recent years (Ostrom 1978; Peyer 2006). All recent phylogenetic analyses have agreed on its placement within Coelurosauria, although its specific relationships have been problematic. It is employed here as a relatively unspecialized coelurosaurian, retaining some of the primitive features of the clade. Peyer (2006) provided a recent diagnosis for C. longipes; although none of these features are autapomorphies, they may represent a unique character combination. Concavenator corcovatus Ortega et al., 2010 2010 Concavenator corcovatus Ortega et al.: 203, figs 1, 3–4. Holotype. MCCM-LH 6666, nearly complete skull and skeleton with soft-tissue impressions. Diagnosis. Allosauroid with: (1) four pneumatic recesses in nasal, three of which are connected; (2) rounded postorbital brow occupying one-third of orbital fenestra; (3) dorsal vertebrae 11–12 with neural spines five times centrum height; (4) caudal vertebrae 2–3 with tall, anteriorly angled neural spines; and (5) pollex terminates at or below distal end of phalanx I of digit II (all from Ortega et al. 2010). Occurrence. Las Hoyas locality, La Cierva township, Cuenca, Spain; Calizas de La Huérguina Formation; late Barremian, Early Cretaceous (Ortega et al. 2010). Remarks. The remarkably well-preserved holotype individual of Concavenator was recently described as a basal carcharodontosaurian, approximately contemporaneous with Neovenator salerii. Although its unusual vertebral morphology superficially resembles that of Becklespinax altispinax (q.v.), Concavenator is quite distinct. Unlike Becklespinax, the posterior dorsal centra bear large lateral fossae (although no foramina), the neural spines have less curved anterior and posterior margins at their bases, and the apices of the tallest spines are anteroposteriorly narrow, curving towards a single apex formed from multiple spines.

2005a Condorraptor currumili Rauhut: 89, figs 2–14. Holotype. MPEF-PV 1672, a left tibia. Hypodigm. Holotype and MPEF-PV 1673–1697, 1700–1705, teeth, cervical, dorsal, sacral and caudal vertebrae, rib fragments, chevron, partial ilium, pubes, ischium, femora, metatarsal IV, and pedal phalanx (probably from the same individual as the holotype). Diagnosis. Megalosauroid theropod with: (1) pleurocoel in anterior cervical vertebrae located immediately posterodorsal to parapophysis; (2) shallow depression on lateral surface of tibia at base of cnemial crest; and (3) metatarsal IV with distinct step dorsally between shaft and distal articular facet (all from Rauhut 2005a). Occurrence. Las Chacritas, 2.3 km west of Cerro Cóndor, Chubut, Argentina; Cañadón Asfalto Formation, Sierra de Olte Group; Bajocian–Callovian, Middle Jurassic (Rauhut 2005a; Volkheimer et al. 2008). Remarks. This Middle Jurassic theropod, penecontemporaneous with Piatnitzkysaurus, was discovered and diagnosed recently (Rauhut 2005a). The fragmentary holotype preserves elements from across the skeleton, providing the means for comparison with other theropods. Its provenance makes it potentially important both evolutionarily and biogeographically. In addition to the holotype, Rauhut (2005a) referred other, disarticulated topotypic remains to Condorraptor on the basis that they all represented a medium-sized theropod with a similar quality of preservation, and thus could belong to a single individual. Most of these remains are closely comparable with those of Piatnitzkysaurus, although they do differ in some respects, suggesting that Condorraptor represents a related but distinct taxon. Rauhut (2005a) described the absence of a posterior incision between the fibular condyle and the medial part of the proximal tibia as an additional autapomorphy. However, the proximal end of MPEF-PV 1672 is highly abraded and the absence of this incision is uncertain. In addition, the presence of a foramen on the lateral surface of the ischial peduncle of the ilium is present variably in other theropods, including Megalosaurus and Piatnitzkysaurus (Benson 2009b), and is not autapomorphic for Condorraptor (contra Rauhut 2005a).

The phylogeny of Tetanurae Cryolophosaurus ellioti Hammer & Hickerson, 1994 1994 Cryolophosaurus ellioti Hammer & Hickerson: 828, figs 2, 3. Holotype. FMNH PR 1821, posterior portion of skull with partial postcranial skeleton.


Diagnosis. Theropod with: (1) transversely oriented, curved midline lacrimal crest bearing fluted anterior and posterior surfaces; (2) lower temporal fenestra constricted by approximated jugal and squamosal; and (3) elongate anterior processes on cervical ribs (all from Smith et al. 2007). Occurrence. Mt. Kirkpatrick, near Beardmore Glacier, central Transantarctic Mountains, Antarctica; Hanson Formation; Sinemurian–Pliensbachian, Early Jurassic (Hammer & Hickerson 1999; Smith et al. 2007). Remarks. Cryolophosaurus is an important taxon for basal theropod studies, as it is represented by a reasonably complete skull and skeleton from a poorly sampled region (Antarctica) during an early interval of theropod evolution. The skull is distinctive, bearing an apomorphic transverse nasolacrimal crest and a nearly bipartite lower temporal fenestra. A recently published study (Smith et al. 2007) has described its morphology and relationships in detail, suggesting that Cryolophosaurus originated just basal to the split between Ceratosauria and Tetanurae. That study also found it to belong to a clade of ‘crested’ theropods more derived than coelophysoids, along with Dilophosaurus, ‘D.’ sinensis (see below) and Dracovenator. Our observations suggest that several putative similarities among these forms are over-split features that all relate to the presence of a cranial crest, and that other skeletal features amongst them are less congruent. Recoding of the characters describing the cranial crest by Brusatte et al. (2010a) resulted in failure to resolve the ‘crested’ clade, but the affinities of most crested taxa remain uncertain.

CV 00214 1983 Szechuanosaurus campi Dong et al.: 56, figs 40–43, pls 18–21. Hypodigm. CV 00214, a partial postcranial skeleton lacking the skull. Occurrence. Wujiaba Quarry, suburbs of Zigong city, Sichuan, China; lower part, Shangshaximiao Formation; Oxfordian–early Kimmeridgian, Late Jurassic (Dong et al. 1983).

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Remarks. This specimen was described in detail and has been exhibited in the Municipal Museum of Chongqing for decades (Dong et al. 1983). Its initial assignment to Szechuanosaurus campi is untenable because (1) the type materials of S. campi (IVPP V.235, V.236, V.238, V.239) consist of teeth, which are non-diagnostic; and (2) no teeth are preserved in CV 00214. A recent restudy of CV 00214 (Chure 2001a) concluded that it represented a new taxon, not yet formally named, into which ‘Szechuanosaurus’ zigongensis (ZDM 9012) should also be subsumed. The published descriptions of CV 00214 show that it exhibits a broad, expansive axial neural spine, quite similar to those of Sinraptor and Yangchuanosaurus. Other aspects of the skeleton demonstrate tetanuran and allosauroid affinities. We retain this specimen as distinct from the type of ‘S.’ zigongensis because we cannot identify autapomorphies shared between them, and the latter derives from the underlying Xiashaximiao Formation. ‘Dilophosaurus’ sinensis Hu, 1993 1993 Dilophosaurus sinensis Hu: 65, fig. 1. Holotype. KMV 8701, a nearly complete skull and skeleton. Hypodigm. Holotype and LDM-LCA 10, a nearly complete skull and skeleton. Diagnosis. Theropod with vertical groove or channel on lateral premaxilla adjacent to contact with maxilla. Occurrence. Qinglongshan near Muchulang Village, Xiyangyi Rural Tribal District, Jinning County, Yunnan, China; lower Lufeng Formation; Hettangian–Sinemurian, Early Jurassic. Remarks. One of the most complete theropods known from the lower Lufeng Formation (Hettangian– Sinemurian), ‘D.’ sinensis has been only preliminarily described (Hu 1993). Although it was originally referred to the genus Dilophosaurus based on the presence of a prominent nasolacrimal crest (Hu 1993), subsequent authors have questioned this assignment (Lamanna et al. 1998). Specifically, several features of the holotype (KMV 8701) differ from the conditions seen in Dilophosaurus, including the number of premaxillary teeth, the shape of the premaxilla and lower temporal fenestra, the extent of the maxillary tooth row and the morphology of the external mandibular fenestra. Indeed, as suggested by Lamanna et al. (1998), the currently understood distribution of cranial crests in theropods undermines their utility as indicators of systematic affinity. Nonetheless, more recent work has supported an affinity between ‘D.’ sinensis, Dilophosaurus and other crested theropods (Smith et al. 2007).


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Certain features hint at a more derived position for ‘D.’ sinensis. The presence of five premaxillary teeth is known elsewhere only in Allosaurus and Neovenator, and the premaxilla is reminiscent of the morphology of tetanurans rather than coelophysoids. In addition, Hu (1993) described the cervicals as opisthocoelous and possessing a single pleurocoel, the scapular blade as long and narrow, and the pubic peduncle of the ilium as larger than the ischial peduncle; these characters are seen in tetanurans and ceratosaurs but not coelophysoids. Finally, the distal ischia are fused, as in Neovenator, Elaphrosaurus and Sinraptor. The two specimens constituting the hypodigm of ‘D.’ sinensis are often confused, thanks largely to the use of LDM LCA 10 (or casts thereof) rather than the holotype in numerous exhibit displays. The two skulls have quite different proportions: KMV 8701 is shorter and taller, with a more abbreviate premaxilla, whereas LDM-LCA 10 is long and low, close to the proportions of D. wetherilli. In addition, the dorsal crests in LDM-LCA 10 appear to end farther posteriorly than those of KMV 8701. However, in spite of these differences there are no distinctions in the coded characters used in our analysis. The two specimens also share an unusual ‘channel’ along the posterior premaxilla adjacent to the maxillary contact, a vertical anterior border of the maxilla (distinct from the condition in Dilophosaurus wetherilli), and a similarly shaped and placed promaxillary fenestra. We suggest that the two specimens represent the products of differential preservation (including distortion), ontogenetic stage or individual growth. Dilophosaurus wetherilli (Welles, 1954) 1954 Megalosaurus wetherilli Welles: 595, pl. 1. 1970 Dilophosaurus wetherilli Welles: 989. Holotype. UCMP 37302, nearly complete subadult skeleton. Hypodigm. Holotype, UCMP 37303, a partial skull and postcranial skeleton, and UCMP 77270, a skull with partial skeleton. Diagnosis. Theropod with: (1) thin, paired nasolacrimal crests extending vertically from skull roof, each with fingerlike posterior projection (modified from Welles 1970); (2) lacrimal with thickened posterodorsal rim (Rauhut 2003); and (3) central ‘cap’ on cervical vertebral neural spines (modified from Welles 1970). Occurrence. Near Tuba City and Rock Head, Coconino County, Navajo Indian Reservation, Arizona, USA; Silty Facies, Kayenta Formation; Early Jurassic (Welles 1954, 1984). Remarks. This well-studied taxon (e.g. Welles 1984) remains the largest known coelophysoid, represented by

several specimens spanning a range of sizes and presumably ontogenetic stages. The nasolacrimal crest, once considered unique among theropods, now appears to have been apomorphic in shape but not presence, as similar structures are also found in distantly related forms. Unspecified minor differences between UCMP 77270 and the other specimens of D. wetherilli caused Welles (1984) to suggest that it belonged to a separate taxon. We note the presence of ‘fused’ maxillary interdental plates in UCMP 37303 compared to the ‘unfused’ interdental plates of UCMP 77270, and the presence of a ‘shelf-like’ trochanteric shelf of the femur in UCMP 77270 versus the mound-like trochanteric region of UCMP 37302. These may represent individual rather than interspecific variations. Tykoski (2005) referred a partial skeleton from Gold Spring, Arizona (TMM 43646) to D. wetherilli. However, direct examination reveals minor differences (e.g. taller maxillary interdental places, presence of a pneumatic fossa on the dorsal surface of the jugal process of the maxilla) and we have not included it in the present hypodigm. Dubreuillosaurus valesdunensis (Allain, 2002) 2002 Poekilopleuron? valesdunensis Allain: 75, figs 2–16. 2005a Dubreuillosaurus valesdunensis Allain: 850, figs 1–11. Holotype. MNHN 1998-13, a nearly complete skull with a fragmentary postcranial skeleton. Diagnosis. Megalosauroid theropod with: (1) ventral notch in posterior outline of braincase between exoccipitalopisthotics and basioccipital (Rauhut 2004); and (2) absence of femoral distal extensor groove (Allain 2005a). Occurrence. Near Conteville, Calvados, BasseNormandie, France; Procerites progracilis Zone, Pierre de Caen, Calcaires de Caen; middle Bathonian, Middle Jurassic (Allain 2002). Remarks. Dubreuillosaurus represents one of the most important European theropod discoveries of the last century, not only because it includes a well-preserved skull, but also because it derives from the Calcaires de Caen and is therefore approximately contemporaneous with Megalosaurus and Poekilopleuron. As such, it has the potential to illuminate the materials assigned to both taxa and clarify their phylogenetic relationships. As noted by Allain (2002, 2005a), Dubreuillosaurus shares numerous synapomorphies with ‘megalosaurs’ such as Eustreptospondylus, Afrovenator and Torvosaurus. Indeed, the maxilla of Dubreuillosaurus is very similar to that of Afrovenator. Dubreuillosaurus shows distinct autapomorphies amongst megalosaurs, but other features listed as diagnostic (Allain 2002, 2005a) are difficult to assess due to lack of comparative material from closely

The phylogeny of Tetanurae related taxa, or their more widespread presence in related forms. The postcranium differs sufficiently from that of Poekilopleuron to warrant inclusion in a distinct genus (Allain 2005a), but there is limited overlap in materials between these two taxa. Unfortunately, the lack of skull material for Poekilopleuron makes further comparisons difficult. Duriavenator hesperis (Waldman, 1974) 1883 Megalosaurus bucklandi Owen: 334, pl. 11. 1974 Megalosaurus hesperis Waldman: 325, pls 42, 43. 2008a Duriavenator hesperis Benson: 58, figs 1, 2.


Holotype. NHMUK R332, the anterior portion of a skull. Diagnosis. Megalosaurid theropod with: (1) deep groove on dorsal surface of jugal process of maxilla containing numerous pneumatic foramina; and (2) array of small foramina in ventral part of maxillary articular surface for premaxilla (all from Benson 2008a). Occurrence. Greenhill, Sherborne, Dorset, England; Garantiana garantiana Subzone, Parkinsonia parkinsoni Zone, upper part of Inferior Oolite Group; late Bajocian, Middle Jurassic (Waldman 1974; Benson 2008a). Remarks. The single known specimen of this taxon includes just the anterior third of a skull, collected in the 19th century (Owen 1883) but only fully prepared years later (Waldman 1974; Benson 2008a). The original specific diagnosis (Waldman 1974, p. 325) relied only on tooth count and dental features to distinguish it from other species of Megalosaurus; these features are now known to occur more widely throughout Theropoda. The skull bones do reveal a few phylogenetically relevant features, comparable to those found in other taxa from the Middle Jurassic of Europe such as Dubreuillosaurus and Eustreptospondylus (Benson 2008a). These include the presence of a distinct ‘kink’ in the maxillary ascending ramus, a paradental groove of the dentary that is wide anteriorly and an enlarged, subcircular third dentary alveolus. Elaphrosaurus bambergi Janensch, 1920 1920 Elaphrosaurus bambergi Janensch: 225, figs 1–5. Holotype. MB.R. (unnumbered), partial skeleton lacking the skull, distal forelimbs, ribs, and distal caudals. Hypodigm. Holotype, MB.R.1755, radius and 1756, distal ischium. Diagnosis. Ceratosaur with: (1) thin ventrolateral lamina bordering posterior cervical pleurocoel ventrally; (2) strongly concave ventral border of cervical vertebrae, with

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apex above mid-height of anterior articular face; (3) scapular blade breadth exceeding height of vertebral column; and (4) extremely wide iliac brevis fossa with nearly horizontal brevis shelf (all from Rauhut 2003). Occurrence. RD, dd, and Dysalotosaurus quarries, Kindope, north of Tendaguru, Mtwara, Tanzania; Middle and ?Upper Dinosaur Members, Tendaguru Formation; late Kimmeridgian–?late Tithonian, Late Jurassic (Janensch 1920, 1925; Aberhan et al. 2002; Schrank 2005). Remarks. This taxon was discussed by Carrano & Sampson (2008), and is currently under detailed restudy (Rauhut & Carrano, in prep.). It has been consistently identified as a ceratosaur in recent analyses (Rauhut 2003; Carrano & Sampson 2008). Eocarcharia dinops Sereno & Brusatte, 2008 2008 Eocarcharia dinops Sereno & Brusatte: 225, figs 1–5. Holotype. MNN GAD2, a postorbital. Hypodigm. Holotype and MNN GAD3–GAD14, skull elements and teeth. Diagnosis. Allosauroid theropod with: (1) enlarged, subtriangular, laterally exposed promaxillary fenestra larger in size than maxillary fenestra; (2) circular accessory pneumatic fenestra on posterodorsal ramus of maxilla; (3) dorsoventral expansion of antorbital fossa ventral to promaxillary and maxillary fenestrae; (4) postorbital brow bearing finely textured ovoid swelling above posterodorsal corner of orbit; (5) postorbital medial process with plate-shaped projection fitted to articular slot on frontal; (6) postorbital articulation for jugal includes narrow, laterally facing facet; (7) enlarged prefrontal lacking ventral process, with subquadrate exposure on dorsal skull roof and within orbit; and (8) low protuberance on frontoparietal suture (all from Sereno & Brusatte 2008). Occurrence. G88 and other sites along the Gadoufaoua outcrop, Agadez, Niger; Elrhaz Formation, Tegama Group; Aptian–Albian?, Early Cretaceous (Sereno & Brusatte 2008). Remarks. Eocarcharia shares several features with carcharodontosaurids but appears to represent a primitive member of that clade. In particular, the maxilla is less heavily sculptured, the postorbital-squamosal suture is planar rather than helical, and the prefrontal remains unfused to the lacrimal (Sereno & Brusatte 2008). Eustreptospondylus oxoniensis Walker, 1964 1890 Megalosaurus bucklandi (sic) Woodward & Sherborn: 249.

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1905 Streptospondylus cuvieri (partim) Nopcsa: 293. 1964 Eustreptospondylus oxoniensis Walker: 124, fig. 17e. 2003 Magnosaurus oxoniensis (Walker); Rauhut: 20. Holotype. OUMNH J.13558, partial skull and skeleton of a juvenile individual.


Hypodigm. Holotype and OUMNH J.29775, a left ilium. Diagnosis. Megalosauroid theropod with: (1) shallow lacrimal fenestra incorporating second, smaller foramen (Sadleir et al. 2008); (2) squamosal with hypertrophied ventrolateral flange obscuring posterodorsal corner of lower temporal fenestra in lateral view (Rauhut 2003); (3) absence of ventral midline ridge on posterior cervical and anterior dorsal vertebral centra (Sadleir et al. 2008); (4) marked depression located anteriorly on ventral surface of tenth presacral vertebra (Sadleir et al. 2008); (5) lateral wall of iliac brevis fossa nearly horizontal, exposing medial wall of fossa along entire length in lateral view; and (6) pubic peduncle of ilium as broad anteroposteriorly as mediolaterally (modified from Benson 2009a). Occurrence. Summertown brick pit, Wolvercot and Littlemore, Oxfordshire, England; Peltoceras athleta Zone, Stewart Member, Oxford Clay Formation and Corallian Formation; late Callovian, Middle Jurassic (Huene 1926a; Walker 1964). Remarks. The holotype specimen of Eustreptospondylus oxoniensis has had a lengthy history in theropod studies, as it long represented the most complete theropod skeleton from Europe. Unfortunately, the taxonomic history of the specimen has been equally long and rather convoluted. Originally assigned to Megalosaurus (Woodward & Sherborn 1890), for many years it was considered the best specimen of Streptospondylus cuvieri Owen, 1842 (a nomen dubium; see below) based on presumed similarities with the material now called Streptospondylus altdorfensis (Nopcsa 1905, 1906; Huene 1926a; Swinton 1955). It was assigned to its own genus by Walker (1964). Although the holotype of Eustreptospondylus is a subadult specimen, its completeness makes it important in deciphering ‘megalosaur’ anatomy and relationships. In addition, much of the skull, including the braincase, is preserved and shows similarities to those of Torvosaurus and Piatnitzkysaurus (Bakker et al. 1992; Sereno et al. 1994; Rauhut 2004). Nopcsa’s (1906) lengthy description was never translated into English and so was not widely cited afterwards; only recently has the taxon been redescribed in appropriate detail (Sadleir et al. 2008). Rauhut (2003) synonymized Eustreptospondylus with Magnosaurus based on the shared presence of a transversely and dorsoventrally expanded anterior end of the dentary, an enlarged third dentary tooth, and a shallow

longitudinal nutrient furrow with a rectangular cross section (Rauhut 2003, p. 20). Although these features are similar, as discussed below (see Magnosaurus) there are also several differences in the detailed morphology of preserved materials (Sadleir et al. 2008; Benson 2010b). Notably also, Magnosaurus is represented by a more mature individual that is approximately the same size as the juvenile holotype of Eustreptospondylus. We do not consider them sufficiently similar to warrant synonymy and treat them as separate taxa here. Fukuiraptor kitadaniensis Azuma & Currie, 2000 2000 Fukuiraptor kitadaniensis Azuma & Currie: 1737, figs 2–18. Holotype. FPDM-V97122, associated cranial and appendicular bones (including V96082443, a left humerus). Hypodigm. Holotype; FPDM-V9712229, fragmentary left maxilla, dentary, teeth, cervical vertebrae, dorsal neural arch, and coracoid; V97082553, left humerus; 97081115, V97082120, V980723005, right humeri; V990410001, manual phalanx I-1; V980801141, V980815162, V9912141, manual unguals; V97080937, V98072302, V99090901, V98120001–0002, left femora; V97122BNA2, V97122BNA12, V970730003, V97081201, V97081330, V970813046, V970821039, V980805018, V9708102884, V980813017, V9812638, right femora; V97081317, V970814001, V970820060, right tibiae; V98082026, pedal phalanx III-2; and 42 isolated teeth (FPDM various nos.; Currie & Azuma 2006). Diagnosis. Avetheropod with: (1) proportionally long forearm (ulna : humerus ratio = 0.92); and (2) pubic peduncle of ilium approximately as broad anteroposteriorly as mediolaterally (all from Azuma & Currie 2000). Occurrence. Kitadani locality, along Sugiyama River, northern part of Katsuyama city, Fukui Prefecture, Japan; Kitadani Formation, Akaiwa Subgroup, Tetori Group; Barremian, Early Cretaceous (Kutoba 2005; Currie & Azuma 2006). Remarks. Fukuiraptor has not been included in many phylogenetic analyses since its original description, but both anatomical descriptions (Azuma & Currie 2000; Currie & Azuma 2006) have placed it as a carnosaur (sensu Rauhut, 2003) outside allosauroids and megalosauroids (spinosauroids). It is not clear whether such a position resulted from the fragmentary nature of the available materials or the mixed phylogenetic signal derived from them. Azuma & Currie (2000) suggested that long oblique ‘blood grooves’ (interdenticular sulci) on the teeth were an autapomorphy of Fukuiraptor but they have since been reported in other theropods, including Megalosaurus (Benson 2009b),

The phylogeny of Tetanurae and can therefore be used as a phylogenetically informative character (Benson 2010a). More recently discovered taxa, including Australovenator (Hocknull et al. 2009) and Aerosteon (Sereno et al. 2008), exhibit similar character combinations. For example, although Fukuiraptor appears to possess an autapomorphically long forearm (relative to the arm), a proportionally long forelimb (relative to the hind limb) is also seen in Chilantaisaurus. Giganotosaurus carolinii Coria & Salgado, 1995 1995 Giganotosaurus carolinii Coria & Salgado: 225, figs 1, 2.


Holotype. MUCPv-Ch 1, partial skeleton. Hypodigm. Holotype and MUCPv-52, 95, a dentary. Diagnosis. Allosauroid with two pneumatic foramina on medial surface of quadrate (Coria & Salgado 1995). Occurrence. South and west of El Chocón, Lake Esquiel Ramos Mexia, Neuquén, Argentina; Candeleros Formation, R´ıo Limay Subgroup, Neuquén Group; ?late Cenomanian, Late Cretaceous (Leanza et al. 2004; Corbella et al. 2004). Remarks. Giganotosaurus has received a great deal of attention because of its enormous size, comparable to that of other carcharodontosaurids and in some ways rivalling the largest tyrannosaurids. However, although the holotype is relatively complete, a full description has not yet been published and only the braincase has been documented in detail (Coria & Currie 2002). The original diagnosis of Giganotosaurus (Coria & Salgado 1995) contains numerous features found in more recently discovered specimens belonging to Acrocanthosaurus (Currie & Carpenter 2000) and Mapusaurus (Coria & Currie 2006), and which therefore can no longer be considered diagnostic. The reconstructed skull (Coria & Salgado 1995) includes a posteriorly oriented quadrate that outlines a trapezoidal lower temporal fenestra. This unusual configuration is genuine but several other skull contacts are not preserved, leading to ambiguity regarding its total length. We believe the original skull reconstruction is likely too long (153 cm), and as with Carcharodontosaurus (see above) we consider Giganotosaurus to have had a skull almost exactly comparable in length to that of Tyrannosaurus. Likewise, our measurements of femur length in the holotype (136.5 cm, left) record a smaller size than originally reported (143 cm; Coria & Salgado 1995) and therefore an animal of lower overall body mass. Irritator challengeri Martill et al., 1996 1996 Irritator challengeri Martill et al.: 5, figs 2–4.

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Holotype. SMNS 58022, a nearly complete skull lacking the tip of the rostrum. Diagnosis. Spinosaurid theropod with nasals exhibiting prominent median bony crest that terminates posteriorly in knob-like, somewhat dorsoventrally flattened projection (Sues et al. 2002). Occurrence. Near Buxéxé, 5 km south of Santana do Cariri, Araripe Basin, southern Ceará, Brazil; Romualdo Member, Santana Formation; Albian, Early Cretaceous. Remarks. Originally described as a maniraptoran theropod (Martill et al. 1996), a more recent redescription has identified Irritator as a spinosaurid (Sues et al. 2002). The slightly disarticulated, subadult holotype skull is the most complete known for any spinosaurid, and provides important information about skull morphology and proportions in this group. It also highlights the differences between baryonychines such as Baryonyx and Suchomimus, and spinosaurines such as Irritator, Angaturama and Spinosaurus. The presence of a broad lateral shelf on the surangular was listed as an autapomorphy of Irritator (Sues et al. 2002), but a similar morphology is also present in Baryonyx. However, the combination of spinosaurine features elsewhere in the skull of Irritator serves to distinguish it from baryonychines. As the surangular is unknown in Suchomimus, it cannot be determined whether this feature is homoplastic within Spinosauridae or a symplesiomorphy within the clade. Leshansaurus qianweiensis Li et al., 2009 2009 Leshansaurus qianweiensis Li et al.: 1203, pls 1–3. Holotype. QW 200701, a partial skull and postcranial skeleton. Diagnosis. Tetanuran theropod with distinct ventral ridge on all sacral vertebral centra (Li et al. 2009). Occurrence. Xiaogu, Qianwei County, Sichuan, China; Shangshaximiao Formation; Late Jurassic. Remarks. Leshansaurus was assigned to Sinraptoridae in its original description (Li et al. 2009). However, the braincase bears distinct similarities to those of Piveteausaurus and Dubreuillosaurus whereas the maxilla resembles those of Afrovenator and Duriavenator. There are no clear synapomorphies with members of Allosauroidea. Therefore Leshansaurus may represent an Asian megalosaurid. Lourinhanosaurus antunesi Mateus, 1998 1998 Lourinhanosaurus antunesi Mateus: 112, figs 1–5.

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Holotype. ML 370, a partial postcranial skeleton. Diagnosis. Tetanuran theropod with: (1) triangular spurs anterior to neural spines of even most proximal caudal vertebrae (modified from Mateus 1998); and (2) medial condyle of tibia half the transverse width of fibular condyle.


Occurrence. Peralta, near Lourinhã, Estremadura, Portugal; Sobral Formation; late Kimmeridgian–early Tithonian, Late Jurassic. Remarks. Lourinhanosaurus was originally described as an allosauroid (Mateus 1998) and subsequently recovered as a metriacanthosaurid by Benson (2010a) and Benson et al. (2010). The skeleton is preserved in articulation but many potentially informative elements are incomplete or poorly preserved, and little systematically informative detail is observable. Magnosaurus nethercombensis (Huene, 1923) 1923 ‘Megalosaurus’ nethercombensis Huene: 450. 1926a Megalosaurus (subgen. b) nethercombensis Huene; Huene: 72. 1932 Magnosaurus nethercombensis (Huene); Huene: 220. Holotype. OUMNH J.12143, fragmentary postcranial skeleton and dentaries. Diagnosis. Megalosauroid theropod with obliquely oriented foramina located ventrally on lateral surface of dentary that are elongated to resemble grooves. Magnosaurus can also be distinguished from all other temporally and geographically proximate theropods (Benson 2010b). Occurrence. Nethercomb, 1.6 km north of Sherbourne, Dorset, England; Stephanoceras humphriesianum Zone and Subzone, middle part of Inferior Oolite; early Bajocian, Middle Jurassic (Huene 1923, 1926a; Waldman 1974). Remarks. Originally described as a species of Megalosaurus and later allocated its own genus, Magnosaurus, this taxon is based on fragmentary remains that have proven difficult to interpret phylogenetically. Waldman (1974) focused on the morphology of the dentaries and associated teeth to support its assignment to Megalosaurus but made few comparisons with other theropods. Rauhut (2003) found several potential synapomorphies of the dentary to support a relationship with Eustreptospondylus. In both cases, affinities with traditional ‘megalosaurs’ were implied, but this was only demonstrated in the context of a cladistic analysis recently (Benson 2010a). A detailed redescription of the holotype, including fragmen-

tary postcranial material, has also been published (Benson 2010b). Majungasaurus crenatissimus (Depéret, 1896) 1896 Megalosaurus crenatissimus Depéret: 188, pl. 4, figs 4–8. 1928 Dryptosaurus crenatissimus (Depéret); Depéret & Savornin: 262. 1955 Majungasaurus crenatissimus (Depéret); Lavocat: 259, fig. 1. 1979 Majungatholus atopus Sues & Taquet: 634, fig. 1. Neotype. MNHN MAJ-1, a left dentary. Hypodigm. Neotype; FSL 92.289, 92.290, 92.306, and 92.343 (syntype series, Megalosaurus crenatissimus Depéret, 1896); MNHN MAJ-4 (holotype, Majungatholus atopus Sues & Taquet, 1979); FMNH PR 2008, 2100; and UA 8678. Diagnosis. Abelisaurid with: (1) thickened, fused, highly pneumatic nasals bearing large, bilateral foramina; (2) thin nasal lamina separating left and right premaxillary nasal processes; (3) maxilla with 17 teeth; (4) frontals with median hornlike projection; and (5) pronounced median fossa on sagittal crest (modified from Sampson et al. 1998; Krause et al. 2007; Sampson & Witmer 2007). Occurrence. Meravana and Berivotra, Mahajanga Basin, Mahajanga, Madagascar; Anembalemba Member, Maevarano Formation; Maastrichtian, Late Cretaceous (Depéret 1896; Thévenin 1907; Sampson et al. 1998). Remarks. The history of this taxon, and of its junior synonym Majungatholus atopus, has been given in detail elsewhere (Sampson et al. 1996, 1998; Krause et al. 2007) Mapusaurus roseae Coria & Currie, 2006 2006 Mapusaurus roseae Coria & Currie: 74, figs 2–34. Holotype. MCF PVPH-108.1, a right nasal. Hypodigm. Holotype and MCF PVPH-108.5, 45, 83, 90, 115, 125, 128, 165, 167, 177, 179, 202, representing most elements of the skull and postcranium. Occurrence. Cañadón del Gato, Cortaderas area, 20 km south-west of Plaza Huincul, Neuquén, Argentina; Huincul Formation, R´ıo Limay Subgroup, Neuquén Group; Turonian–Santonian, Late Cretaceous (Dingus et al. 2000; Corbella et al. 2004; Leanza et al. 2004). Remarks. The third carcharodontosaurid described from Argentina, Mapusaurus is known from a bonebed deposit containing several disarticulated individuals, some of which


The phylogeny of Tetanurae approximated Giganotosaurus in size. Coria & Currie (2006) united Mapusaurus and Giganotosaurus as sister taxa within the subfamily Giganotosaurinae. It is difficult to reliably distinguish elements of Mapusaurus from their counterparts in Giganotosaurus independent of their stratigraphical origin. Several of the autapomorphies provided for Mapusaurus (Coria & Currie 2006) are not observable in Giganotosaurus, while others must be reinterpreted. It appears to differ from Giganotosaurus in lacking a second pneumatic foramen on the medial quadrate and in details of the topology of the nasal rugosities. We cannot confirm any of the autapomorphies of Mapusaurus proposed by Coria & Currie (2006). Several cannot be observed in Giganotosaurus due to incomplete preservation, including the presence of an upper quadratojugal process of jugal split into two prongs; a small anterior mylohyoid foramen positioned above dentary contact with splenial; and a humerus with broad distal end and little separation between condyles (Coria & Currie 2006). The supposed fusion between metacarpals II and III is based on a specimen (PVPH 108-48) that we consider to be a distal humerus. As figured (Coria & Currie 2006, fig. 26), the ilium of Mapusaurus is long and low relative to that of Giganotosaurus. However, this bone is poorly preserved and its outer margins represent broken surfaces, so the original shape is unclear. Marshosaurus bicentesimus Madsen, 1976b 1976b Marshosaurus bicentesimus Madsen: 51, figs 1–5. Holotype. UMNH VP 6373 (= UUVP 2826), a left ilium. Hypodigm. Holotype and UMNH VP 7820 (= UUVP 3266), right premaxilla; 7824 (= UUVP 1846), right maxilla; (= UUVP 1864), left maxilla; 7825 (= UUVP 4695), right maxilla; 6364 (= UUVP 40-555), left dentary; 6367 (= UUVP 3454), left dentary; 6368 (= UUVP 3502), right dentary; 6372 (= UUVP 1845), left ilium; (= UUVP 1182), right ilium; 6374 (= UUVP 2742), right ilium; 6384 (= UUVP 40-295), left pubis; 6386 (= UUVP 1867), left pubis; 6387 (= UUVP 4736), right pubis; 6379 (= UUVP 2832), right ischium; 6380 (= UUVP 2878), left ischium (paratypes); CMNH 21704 (= DINO 16455b, DMN 343), partial skull, vertebrae, scapula and forelimb; DMNH 3718, partial skull and vertebrae; YPM-PU 72-1, partial pelvis. Diagnosis. Megalosauroid theropod with: (1) no prominent ventral keels on posterior cervical and anterior dorsal vertebrae; and (2) articular surface of pubic peduncle divided into anterior bulge and posterior concavity (modified from Madsen 1976b). Occurrence. Cleveland-Lloyd Dinosaur Quarry, Emery County and Dinosaur National Monument, Uintah County,

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Utah; DMNH loc. 882, Moffat County and Dry Mesa Quarry, Montrose County, Colorado, USA; Brushy Basin Member, Morrison Formation; Kimmeridgian, Late Jurassic. Remarks. Marshosaurus was originally described from dissociated remains found at the Cleveland-Lloyd Quarry in Utah (Madsen 1976b), a few years after another enigmatic theropod, Stokesosaurus, was also identified from the same deposit (Madsen 1974). For years thereafter, isolated medium-sized theropod bones from the Morrison Formation could only be assigned preliminarily to one or the other taxon (e.g. Britt 1991). Recent discoveries have improved this situation. A new specimen of Marshosaurus from Dinosaur National Monument (CMNH 21704) provides confirmation of the original associations of cranial materials proposed by Madsen (1976b). A partial skeleton of Stokesosaurus was also recently reported from England (Benson 2008b) that permits postcrania to be more reliably assigned. However, we must still base some associations on the knowledge that Marshosaurus is a relatively primitive tetanuran whereas Stokesosaurus is a tyrannosauroid. These can only be confirmed by the discovery of a partial skeleton that includes an ilium similar to the holotype. Although similar to both Condorraptor and Piatnitzkysaurus, Marshosaurus differs from both in exhibiting gently convex (rather than flat) anterior centrum faces in the anterior and middle cervicals; this is likely plesiomorphic. Madsen (1976b, fig. 3A) described an imperforate maxillary fenestra divided into two pockets instead of the usual single pocket as an autapomorphy of M. bicentesimus based on a referred left maxilla (UMNH VP 7825). However, this feature is difficult to identify in recently referred materials (CMNH 21704) and may vary between individuals. The pubis (UMNH VP 6387) shows a small anterior bulge, absent in other megalosauroids, that forms a blunt ‘peg-and-socket’ articulation with the ilium. This is the reverse of the condition seen in ceratosaurs, in which the ilium bears a ‘peg’ that fits into a ‘socket’ on the pubis (Carrano et al. 2002). Marshosaurus is more completely known than either Condorraptor or Piatnitzkysaurus and includes better-preserved maxillae and braincases that share derived features with megalosaurs. Masiakasaurus knopfleri Sampson et al., 2001 2001 Masiakasaurus knopfleri Sampson et al.: 504, figs 1, 2. Holotype. UA 8680, a left dentary. Hypodigm. Holotype and numerous additional specimens listed by Carrano et al. (2011). Diagnosis. Abelisauroid with: (1) four anteriormost dentary teeth procumbent, with first set in large, ventrally expanded alveolus that is almost horizontally oriented;

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and (2) strongly heterodont lower dentition, grading from elongate, weakly serrated, apically round teeth with labiolingually positioned carinae (anteriorly) to increasingly recurved, transversely compressed teeth with mesiodistally positioned carinae (posteriorly) (all from Sampson et al. 2001).


Occurrence. Berivotra, Mahajanga Basin, Mahajanga, Madagascar; Anembalemba Member, Maevarano Formation; Maastrichtian, Late Cretaceous (Carrano et al. 2002). Remarks. This taxon has been discussed in detail elsewhere (Carrano et al. 2002, 2011), including new materials that clarify previously unknown aspects of its anatomy (Carrano et al. 2011). More than two-thirds of the skeleton is now known. Megalosaurus bucklandii Mantell, 1827 1824 Megalosaurus Buckland: 391, pls 40–47. 1826 Megalosaurus conybeari von Ritgen: 354 (nomen nudum). 1827 Megalosaurus bucklandii Mantell: 67, pl. 18, fig. 2; pl. 19, figs 1, 8, 12, 14–16. 1832 Megalosaurus bucklandi Mantell; von Meyer: 110 (lapsus calami). Lectotype. OUMNH J.13506; partial right dentary. Hypodigm. Lectotype and numerous additional specimens listed by Benson (2010a). Diagnosis. Megalosauroid theropod with a unique combination of: (1) 13–14 dentary teeth (modified from Waldman 1974); (2) unexpanded third dentary alveolus (Benson 2010a); (3) dentary straight in dorsal view with unexpanded symphyseal area (modified from Waldman 1974); (4) lateral row of neurovascular foramina housed in shallow longitudinal groove in lateral dentary surface (Benson 2010a); (5) tall, unfused paradental plates on dentary (Benson 2010a); (6) two Meckelian foramina (Benson 2010a); and (7) shallow Meckelian groove (Benson 2010a). The referred materials of M. bucklandii exhibit additional autapomorphies, including: (1) evenly rounded ventral surfaces of sacral centra 1 and 3–5, with angular, longitudinal ridge on ventral surface of sacral centrum 2 (Benson 2010a); (2) dorsally directed flange at mid-height of scapular blade (Waldman, 1974); (3) array of posterodorsally inclined grooves on lateral surface of median iliac ridge (Benson 2010a); (4) anteroposteriorly thick ischial apron with nearly flat medial surface (Benson 2010a); (5) prominent, rugose distal ischial tubercle (Benson 2010a); and (6) complementary groove-and-ridge structures on articular surfaces between metatarsals II and III (Benson 2010a).

Finally, M. bucklandii possesses several characters that are not unique but are absent in all other megalosaurids, such as: (1) pneumatic jugal; (2) unexpanded third dentary alveolus; (3) dorsal neural spines more than 1.9 times centrum height; and (4) anterolaterally inclined deltopectoral crest. Uniquely among megalosauroids, the maxillary anterior process is taller dorsoventrally than it is long anteroposteriorly (Benson 2010a). Occurrence. Stonesfield, near Woodstock, 19 km northwest of Oxford, and Sarsgrove and Workhouse Quarry, Chipping Norton, Oxfordshire; New Park Quarry and Oakham Quarry, Gloucestershire, England; Stonesfield Slate, Taynton Limestone Formation, Chipping Norton Limestone Formation and Sharp’s Hill Formation; lowest middle Bathonian, Middle Jurassic. Remarks. The first dinosaur formally described (Buckland 1824), M. bucklandii has remained problematic ever since. Known from disassociated, and probably isolated, bones deposited allochthonously in the marine Stonesfield Slate facies of the Taynton Limestone Formation, the type series of M. bucklandii was long suspected to comprise at least two distinct theropod taxa (Allain & Chure 2002; Day & Barrett 2004). Without definitive associations of elements, it has been difficult to determine which materials belong to each ‘taxon’. In addition, for decades it was commonplace to ascribe to the genus Megalosaurus theropod specimens derived from Late Triassic through Late Cretaceous strata worldwide, resulting in at least 50 nominal taxa (Benson 2009a). The consequence of this was a diluted perception of the true morphology of the original materials, and the emergence of widespread opinion that Megalosaurus was a truly generalized theropod in almost all respects. In subsequent years new genera were named for most of the diagnostic species referred to Megalosaurus, but only recently has M. bucklandii been adequately restudied. This is unfortunate because aside from the historical interest in M. bucklandii, many of the Stonesfield Slate theropod specimens resemble those of Torvosaurus (Galton & Jensen 1979; Britt 1991) and Eustreptospondylus (Bakker et al. 1992) and therefore may represent an early member of their lineage. These materials suggest that at least one Stonesfield Slate taxon was not ‘generalized’ but exhibited a morphology characteristic of one particular theropod clade. New research has clarified these materials and the morphology of M. bucklandii (Benson et al. 2008; Benson 2009b, 2010a). As a result, we can be much more confident that the majority (perhaps all) large theropod specimens from the Stonesfield Slate pertain to a single taxon. In addition, we have further justification in recognizing M. bucklandii as the only valid, correctly attributed species of the genus Megalosaurus. Several autapomorphies of the

The phylogeny of Tetanurae topotype material allow referral of specimens from other British Bathonian localities (Benson 2010a). Megaraptor namunhuaiquii Novas, 1998 1998 Megaraptor namunhuaiquii Novas: 4, figs 1–3. Holotype. MCF-PVPH 79, a right ulna, left manual phalanges I-1 and I-2, and distal right metatarsal III.


Hypodigm. Type and MUCPv 341, a partial postcranial skeleton. Diagnosis. Tetanuran with: (1) blade-like olecranon process of ulna (Novas 1998); (2) manual phalanx I-1 subtriangular in proximal view with dorsal portion wider than ventral (Novas 1998); and (3) distal end of metatarsal IV shaft narrower than midshaft (Calvo et al. 2004). Occurrence. Sierra de Portezuelo and Futalognko Quarry, Lago Barreales, Neuquén, Argentina; Portezuelo Formation, Neuquén Group; late Turonian–middle Coniacian, Late Cretaceous (Novas 1998; Calvo et al. 2004). Remarks. Megaraptor remains a poorly known theropod, originally founded on very incomplete but intriguing materials that initially suggested coelurosaurian affinities (Novas 1998). More recent discoveries in the Futalognko Quarry have provided additional anatomical information that clarifies its phylogenetic position (Calvo et al. 2002, 2004; Porfiri & Calvo 2003). The enlarged ungual, originally ascribed to the pes, has been shown to belong to manual digit I. In addition, the remainder of the skeleton shows similarities to carcharodontosaurians and other basal tetanurans, rather than coelurosaurs. Calvo et al. (2004) noted that carcharodontosaurid teeth have been found in the same quarry as the MUCPv 341 but that in the absence of articulated specimens they could not be referred unequivocally to Megaraptor. However, one of the revised diagnostic characters of Megaraptor, an elongate anterior pleurocoel in the mid-cervical vertebrae (Calvo et al. 2004, p. 567), is also present in Giganotosaurus, Neovenator, some specimens of Allosaurus, and a mid-cervical vertebra from Morocco assigned to Carcharodontosaurus saharicus (UCPC OT6). Similarly, another putative autapomorphy, prominent anterior and posterior centrodiapophyseal laminae on proximal caudal vertebrae, can also be observed in Aerosteon and Orkoraptor.

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Diagnosis. Allosauroid with ventral surfaces of posterior dorsal centra flat and with breadth approximately 2/3 posterior height of centrum (Benson 2009a). Occurrence. Jordan’s Cliff, Overcombe, 1.5 km north of Weymouth, Dorset, England; Cardioceras cordatus Zone, Weymouth Member, Upper Oxford Clay; early Oxfordian, Late Jurassic (Huene 1926a, Walker 1964). Remarks. Originally described as a species of Megalosaurus, Metriacanthosaurus was removed to its own genus by Walker (1964), who noted the unusual shape of the dorsal margin of the ilium, moderate elongation of the dorsal vertebral neural spines, and fused distal ischia and pubes. Unfortunately, the dorsal margin of the ilium is sufficiently damaged and incomplete that the outline proposed by Walker (1964, fig. 16d) cannot be confirmed and is here regarded as doubtful. Nonetheless, the morphologies of the ilium, ischium and dorsal vertebrae are distinct enough from those of Megalosaurus to warrant retention of Metriacanthosaurus as a separate taxon. It also occurs considerably later in time than M. bucklandii and shows various similarities with penecontemporaneous Chinese ‘sinraptorids’ (Paul 1988a; Benson 2009a). Monolophosaurus jiangi Zhao & Currie, 1994 1992 Monolophosaurus jiangjunmiaoi Dong.: 71 (nomen nudum). 1994 Monolophosaurus jiangi Zhao & Currie: 2028, figs 1–5. Holotype. IVPP 84019, a complete skull and skeleton. Diagnosis. Tetanuran with: (1) nasal process of premaxilla bifurcated posteriorly (Brusatte et al. 2010a); (2) lateral surface of premaxilla with deep groove between subnarial foramen and foramen on base of nasal process (Brusatte et al. 2010a); (3) large midline crest formed by nasals with straight dorsal margin nearly parallel to maxillary alveolar margin (modified from Zhao & Currie 1994); (4) two enlarged, subequal pneumatic fenestrae in posterodorsal part of narial fossa (Brusatte et al. 2010a); (5) lacrimal with discrete, tab-like process projecting dorsally above preorbital bar (Brusatte et al. 2010a); and (6) rectangular frontals, much wider than long (width/length = 1.67) (Brusatte et al. 2010a).

1923 Megalosaurus parkeri Huene: 453. 1964 Metriacanthosaurus parkeri (Huene); Walker: 109, fig. 16.

Occurrence. 34 km north-east of Jianjungmiao, Junggar Basin, Xinjiang Uygur Zizhiqu, China; lower Shishugou (= Wucaiwan) Formation; middle Bathonian–late Callovian, Middle Jurassic (Zhao & Currie 1994; Eberth in Brusatte et al. 2010a and Zhao et al. 2010).

Holotype. OUMNH J.12144, a partial postcranial skeleton.

Remarks. One of the few well-represented Middle Jurassic theropods, Monolophosaurus was originally described

Metriacanthosaurus parkeri (Huene, 1923)


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as a megalosaurid, though this was explicitly considered as an evolutionary grade (Zhao & Currie 1994). Its distinctive skull has drawn much attention, but the holotype also includes most of the axial column and some appendicular elements. Most phylogenetic analyses based on the original, brief description have recovered Monolophosaurus as an allosauroid (Sereno et al. 1996; Rauhut 2003; Holtz et al. 2004), with which it shares several pneumatic features of the skull. However, Smith et al. (2007) noted some comparatively primitive features and recovered Monolophosaurus as a basal avetheropod, and Benson (2010a) found it to be a megalosauroid closely related to Chuandongocoelurus. Recent detailed descriptions of the holotype noted numerous unusual and apparently primitive features of the skeleton, and the affinities of Monolophosaurus remain far from certain (Zhao et al. 2010; Brusatte et al. 2010a). Neovenator salerii Hutt et al., 1996 1996 Neovenator salerii Hutt et al.: 638, figs 2–4. Holotype. NHMUK R10001 and MIWG 6348, partial skull and skeleton divided between the two institutions. Hypodigm. Holotype; MIWG 5470, two vertebrae and 6352, fragmentary postcranial skeleton. Diagnosis. Allosauroid theropod with: (1) accessory interpremaxillary peg-and-socket articulation in dorsal part of premaxillary symphysis (Brusatte et al. 2008); (2) large maxillary fenestra approximately 1/6 length of maxillary tooth row (modified from Hutt et al. 1996); (3) transverse expansion of anterior face of axial intercentrum (Brusatte et al. 2008); (4) lateral foramina on anterior surface of odontoid (Brusatte et al. 2008); (5) ischial distal boot conjoined anteriorly but divergent posterolaterally (Brusatte et al. 2008); (6) femoral head both oriented anteromedially and inclined proximally (Brusatte et al. 2008); (7) robust ridge on external surface of femoral lesser trochanter (Brusatte et al. 2008); (8) thumbprint-shaped depression on posterior surface of femoral shaft, lateral to proximal end of fourth trochanter (Brusatte et al. 2008); (9) proximodistally short, notch-like extensor groove and nearly flat anterior surface of distal femur (Brusatte et al. 2008); and (10) suboval rugosity on medial surface of distal tibia (Brusatte et al. 2008). Relative to other allosauroids, Neovenator is autapomorphic in possessing fusion of the cervical ribs to the posterior cervical vertebrae. In fact, the presence of camellate internal texture exposed externally on the parapophyseal facets of the eighth and ninth cervical vertebrae may record the fusion process of these elements.

Remarks. The holotype of Neovenator is the most completely preserved theropod from the Wealden Group of England. It was the first definite allosauroid recognized from Europe (although others have since been described; Pérez-Moreno et al. 1999; Rauhut & Fechner 2005; Mateus et al. 2006; Ortega et al. 2010) and has been considered the latest surviving allosaurid (Hutt et al. 1996; Smith et al. 2007). Since its original description, the placement of Neovenator within Allosauroidea has varied, with some authors (Holtz et al. 2004; Brusatte & Sereno 2008; Benson 2010a; Benson et al. 2010) favouring a closer relationship to Carcharodontosauridae than Allosauridae. Recent restudy of Neovenator has revealed a suite of distinctive autapomorphies, along with numerous synapomorphies of carcharodontosaurians (Naish et al. 2001; Brusatte et al. 2008). Many of the autapomorphies proposed by Brusatte et al. (2008) are also present in the recently described Aerosteon and Australovenator (Benson et al. 2010), and others may also prove to be more widely distributed once more complete materials are known. Ornitholestes hermanni Osborn, 1903 1903 Ornitholestes hermanni Osborn: 459, figs 1–3. 1970 Coelurus hermanni (Osborn); Steel: 15. Holotype. AMNH 619, nearly complete skull and partial skeleton. Diagnosis. Coelurosaurian with: (1) teeth of premaxilla prominent, larger than maxillary teeth and bearing flattened apex; and (2) retroarticular process offset medially from lateral edge of mandible (all from Rauhut 2003). Occurrence. Bone Cabin Quarry, near Medicine Bow, Albany County, Wyoming, USA; Morrison Formation; Kimmeridgian–Tithonian, Late Jurassic. Remarks. This small theropod was one of the earliest wellknown coelurosaurs from the Jurassic. Although known from a nearly complete skeleton (Osborn 1903, 1917), Ornitholestes apparently lacks many of the specializations that characterize most individual coelurosaur clades and as a result, like Compsognathus, it has held many phylogenetic positions within Coelurosauria. For the purposes of this study, it is used as a representative coelurosaur because it retains many plesiomorphic character states of the clade. Piatnitzkysaurus floresi Bonaparte, 1979 1979 Piatnitzkysaurus floresi Bonaparte: 1378, fig. 1. Holotype. PVL 4073, partial skull and skeleton.

Occurrence. Cliffs near Grange Chine, south-west coast of Isle of Wight, England; Wessex Formation; late Hauterivian–early Barremian, Early Cretaceous.

Hypodigm. Holotype and MACN-CH 895, partial skull and skeleton.

The phylogeny of Tetanurae Diagnosis. Tetanuran with: (1) strongly inflated base of maxillary ascending process (Rauhut 2003); and (2) evenly rounded ventral surfaces of most sacral centra, except sacral 3 bears flat midline strip and sacral 5 is broad and flat (Benson 2010a).


Occurrence. Cerro Cóndor, 1.5 km west of the former Farias store, Cerro Cóndor village, right bank of the river, Chubut, Argentina; Cañadon Asfalto Formation, Sierra de Olte Group; Bajocian–Callovian, Middle Jurassic (Bonaparte 1979; Volkheimer et al. 2008). Remarks. Initially assigned to Allosauridae (Bonaparte 1979, 1986; Sereno et al. 1996), Piatnitzkysaurus represents a temporally and geographically important taxon. Today it remains one of the few well-represented theropods from the Middle Jurassic and one of the only diagnosable theropods from the Jurassic of South America. The two known specimens were found in close association along with materials of the sauropods Volkheimeria and Patagosaurus (Bonaparte 1979, 1986). Restudy of the hypodigm of Piatnitzkysaurus reveals that it lacks all the proposed synapomorphies of the Allosauridae and Allosauroidea, instead presenting a set of features characteristic of more basal tetanurans. For example, the pubis has an enclosed obturator foramen and both the pubis and ischium show only modest distal expansions. The maxilla is quite similar to that of Marshosaurus in lacking an anterior ramus and bearing a curved ventral margin, a ventrally extensive antorbital fossa, and two prominent rows of foramina parallel to the tooth row on the lateral surface. Although Piatnitzkysaurus is very similar to Condorraptor, it can be distinguished from this form on the basis of the sacral vertebral centra (in Condorraptor the second sacral centrum is broad and flat, whereas the third is broad and gently concave mediolaterally; Rauhut 2005a). Likewise, Marshosaurus has a straight dentary whereas that of Piatnitzkysaurus curves anteromesially. Piveteausaurus divesensis (Walker, 1964) 1923 Streptospondylus cuvieri Piveteau: 7, pl. 1, figs 1, 2, pl. 2. 1964 Eustreptospondylus divesensis Walker: 124, fig. 17f. 1977 Piveteausaurus divesensis (Walker); Taquet & Welles: 192, figs 1–5. 1988a Proceratosaurus divesensis (Walker); Paul: 304. Holotype. MNHN 1920-7, a braincase. Diagnosis. The limited materials of Piveteausaurus are insufficient to determine autapomorphies for this taxon given that the braincase of Leshansaurus is so strikingly similar. However, based on its provenance and morphology the only reasonable candidate for synonymy

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is Eustreptospondylus, from which it has been specifically distinguished (Walker 1964; see below). Occurrence. Vaches Noires cliffs, near Dives, Calvados, Basse-Normandie, France; Marnes de Dives; late Callovian, Middle Jurassic. Remarks. The holotype of Piveteausaurus, a braincase, was discovered in the Vaches Noires cliffs at Dives and originally described as a specimen of Streptospondylus cuvieri (Piveteau 1923). Walker (1964) made it the type of a second species of Eustreptospondylus, E. divesensis, but it was later removed to its own genus (Taquet & Welles 1977). The braincase is large and weathered, making identification of certain sutures difficult. For example, we could not confirm the exact boundaries of the supraoccipital and exoccipital near the foramen magnum, which were illustrated by Taquet & Welles (1977). Some workers have suggested that Piveteausaurus might be synonymous with Megalosaurus (Buffetaut et al. 1991a; Buffetaut & Enos 1992), but the lack of braincase material from the latter taxon makes this speculative, especially given the stratigraphical distance between these forms. Nonetheless, the relatively anteroposteriorly long parietals and frontals are more similar to those of nonavetheropod tetanurans than allosauroids. The absence of adult braincase specimens from other large megalosauroids prevents determining whether Piveteausaurus truly represents a distinct taxon. The general proportions are dorsoventrally tall and anteroposteriorly narrow, resembling those of large bodied members of other clades (e.g. Allosaurus; Madsen 1976a), but unlike the comparatively long and low braincases of the juvenile holotype materials of Dubreuillosaurus (Allain 2002) and Eustreptospondylus (Sadleir et al. 2008). However, the detailed morphology reveals megalosauroid affinities. For instance, Piveteausaurus possesses a broad fossa on the basioccipital apron ventral to the occipital condyle and more than two-thirds its width, as do spinosaurs and some other megalosaurs (e.g. Dubreuillosaurus, Eustreptospondylus). It also exhibits laterally, rather than ventrolaterally, directed paraoccipital processes, resembling the condition in megalosaurs such as Dubreuillosaurus and Eustreptospondylus, as well as non-tetanuran theropods. Poekilopleuron bucklandii Eudes-Deslongchamps, 1837 1837 Poekilopleuron bucklandii Eudes-Deslongchamps: 79, pls 2, 4–8. 1841 Poikilopleuron bucklandi Eudes-Deslongchamps; Owen: 458 (lapsus calami). 1843 Poecilopleuron bucklandi Eudes-Deslongchamps; Fitzinger: 61(lapsus calami). 1879 Megalosaurus bucklandi (Eudes-Deslongchamps); Hulke: 233. 1923 Megalosaurus poikilopleuron (EudesDeslongchamps); Huene: 451.

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Holotype. Musée du Caen collection (destroyed), a partial postcranial skeleton including caudal vertebrae, ribs, gastralia, forelimb, and hind limb. Hypodigm. Illustrations of type materials (EudesDeslongchamps 1837); MNHN 1897-2, casts of gastralia, humerus, ulna, radius, metacarpals, manual phalanges, pedal phalanges; and YPM 4938, casts of humerus, ulna, and radius.


Diagnosis. Non-coelurosaur tetanuran with ulnar olecranon process absent (Allain & Chure 2002). Occurrence. Quarry on Basse-rue, La Maladrerie, Caen, Basse-Normandie, France; Procerites progracilis Zone, Banc Royal, Calcaire de Caen; middle Bathonian, Middle Jurassic. Remarks. The unfortunate demise of the holotype and only known specimen of Poekilopleuron during World War II has been amply documented (e.g. Bigot 1945; Allain & Chure 2002) and has contributed significantly to the ambiguity surrounding the affinities of this taxon. The specimen originally comprised a partial postcranial skeleton with several phylogenetically informative elements, such as the astragalus, tibia, and forelimb. Although the circumstances of its original discovery and acquisition (EudesDeslongchamps 1837) have lent doubt as to whether the elements were associated, the published details of the account suggest that most or all of the materials pertained to a single individual. Since its discovery, the geographical and geological proximity of Poekilopleuron with Megalosaurus bucklandii has led many researchers to speculate that the two taxa might be synonymous, or that Poekilopleuron might represent a distinct species of Megalosaurus (formally renamed M. poikilopleuron by Huene (1923) to avoid homonymy with the type species). A recent review of Poekilopleuron (Allain & Chure 2002) supported the separation of these taxa, but partly because the authors considered M. bucklandii to be a nomen dubium. They tentatively assigned Poekilopleuron to the Spinosauroidea (i.e. Megalosauroidea), citing a single synapomorphy (the relative length of the deltopectoral crest) and the absence of other features claimed to be typical of more derived theropods. However, it is not clear that all of these features are absent in Poekilopleuron (Benson 2010a). For example, the distalmost preserved chevrons, representing middle caudal elements, show a partial bend or kink, less than that seen in most allosauroids and coelurosaurs but more than in many more primitive theropods (e.g. coelophysoids and ceratosaurs). The long axis of the humeral distal condyles form an oblique angle with the longitudinal shaft axis, rather than a right angle, giving an ‘S’-shaped outline in lateral view unlike the condition in megalosaurids but simi-

lar to that in Allosaurus. The humerus also shows significant obliquity between the long axes of the proximal and distal condyles, which can be observed despite the missing proximal head. Finally, the midshaft median tuberosity on the radial shaft resembles to a similar structure in Acrocanthosaurus (NCSM 14345; Currie & Carpenter 2000). The genus Poekilopleuron (frequently misspelt Poicilopleuron or Poikilopleuron) was often used in the 19th century to receive incomplete specimens of large theropods. These include Poicilopleuron minor Cope, 1878 (indeterminate), Poikilopleuron pusillus Owen, 1876 (now Aristosuchus pusillus; Seeley, 1887), Poekilopleuron schmidti Kirprianow, 1883 (see Fragmentary Taxa, below) and Poicilopleuron valens Leidy, 1870 (now Antrodemus valens; Leidy, 1873). None of these materials pertain to Poekilopleuron, which should be restricted to the type species. Proceratosaurus bradleyi (Woodward, 1910) 1910 Megalosaurus bradleyi Woodward: 114, pl. 13. 1926a Proceratosaurus bradleyi (Woodward); Huene: 69, fig. 40. Holotype. NHMUK R4860, a partial skull missing most of the dorsal surface. Diagnosis. Coelurosaurian with: (1) anterior nasal process of premaxilla inclined slightly anterodorsally, with nasal horn core overhanging premaxillary internarial bar anteriorly; (2) premaxillary internarial bar bifurcated posteriorly; (3) promaxillary foramen located posterodorsal to anterior end of antorbital fossa; and (4) anteriormost dentary tooth curved anteriorly, with carinae oriented labiolingually (all from Rauhut et al. 2010). Occurrence. Minchinhampton Reservoir, Gloucestershire, England; White Limestone Formation, Great Oolite Group; middle–late Bathonian, Middle Jurassic. Remarks. P. bradleyi is represented by most of a skull that was originally described as a species of Megalosaurus (Woodward 1910). Huene (1926a) later recognized its generic distinctiveness and highlighted the unusual structure of the rostrum, which includes the base of a midline horn or crest; this led him to assign the taxon to Ceratosauria. More recently, Proceratosaurus has been consistently assigned to Coelurosauria (Holtz 2000; Rauhut 2003; Holtz et al. 2004), and more specifically has been considered a primitive tyrannosauroid (Rauhut et al. 2010). It is used here to represent basal coelurosaurian character states. Saurophaganax maximus Stovall, 1941 1941 Saurophagus maximus Stovall in Ray: 38 (nomen nudum).

The phylogeny of Tetanurae 1988a Allosaurus amplexus Paul: 312. 1995 Saurophaganax maximus (Stovall); Chure: 106, figs 1, 2. 1998 Allosaurus maximus (Stovall); Smith: 126.


Holotype. OMNH 01123, an anterior dorsal vertebral neural arch. Hypodigm. Holotype and OMNH 01771, postorbital; 01142, 02145, quadrates; 01152, 01153, 01680, teeth; 01135, atlas; 01444, 01446, 02146, 02147, cervical vertebrae; 01450, 01906, anterior dorsal vertebrae; 01433, 01947, sacral vertebrae and ribs; 01122, 01904, 01927, 01928, 10357, proximal caudal vertebrae; 01102, 01104, 01180, 01438, 01439, 01684, 01685, chevrons; 02154, 04016, scapulae; 01693, 01935, humeri; 01364, 01415, radii; 01434, ulna; 01929, metacarpal II; 01127, 01128, 01920, 01921, manual phalanges; 01338, ilium; 01425, 01707, pubes; 01703, 01737, ischia; 01708, 02114, 10381, femora; 01370, 02149, 04666, tibiae; 01681, metatarsal I; 01461, metatarsal II; 01191, 01192, 01924, metatarsal III; 01193, 01306, 01396, metatarsal IV; 01126, 01911, 01912, 01914–01916, 01918, 01919, 01925, 10373, 10375–10377, 10732, 52384–52395, pedal phalanges. Diagnosis. Allosauroid with: (1) atlas lacking prezygapophyses for proatlas; and (2) accessory horizontal lamina at base of dorsal vertebral neural spines, located dorsal and parallel to transverse process (all from Chure 1995). Occurrence. Stovall Quarry 1, east of Kenton, Cimarron County, Oklahoma, USA; upper Brushy Basin Member equivalent, Morrison Formation; ?Tithonian, Late Jurassic (Stovall 1938; Chure 1995).

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Lectotype. IVPP V.2885.1–2885.2, a well-preserved braincase and posterior part of the skull roof. Hypodigm. Lectotype and paralectotype series: IVPP V.2885.3–V.2885.7, partial skull, axis and caudal vertebrae that likely pertain to the same individual as the lectotype (Hu 1964; Brusatte et al. 2009a). Diagnosis. Allosauroid with: (1) maxillary antorbital fossa reduced in extent, nearly absent; (2) paradental groove on medial surface of maxilla absent; (3) deep, dorsoventrally oriented grooves located dorsally on maxillary paradental plates; (4) pneumatic recess penetrates to posterior end of nasal; (5) sagittal crest on frontal; and (6) large pneumatic foramen at anterodorsal corner of dorsal tympanic recess of prootic (all from Brusatte et al. 2009a, 2010b). Occurrence. Maortu, 60 km north of Chilantai (Jilantai), eastern Alashan Desert, Nei Mongol Zizhiqu, China; Ulansuhai Formation; Turonian or younger (< 92 Ma), Late Cretaceous (Kobayashi & Lü 2003). Remarks. The second named species of Chilantaisaurus, C. maortuensis, is represented by a very partial skeleton. The distinct morphologies of these species have led to confusion regarding the affinities of Chilantaisaurus, which has been placed amongst both allosauroids (Hu 1964) and megalosauroids (‘spinosauroids’; Sereno et al. 1998). More recently, the two species of Chilantaisaurus have been found to belong to distinct genera (Chure 2001a), with ‘C.’ maortuensis placed in the new genus Shaochilong which may represent a derived carcharodontosaurid (Brusatte et al. 2009a, 2010b). Shidaisaurus jinae Wu et al., 2009 2009 Shidaisaurus jinae Wu et al.: 9, figs 2–9.

Remarks. The validity of Saurophaganax has been debated since its discovery. Originally named Saurophagus maximus (Ray 1941; a nomen nudum: Chure 1995), the type materials derived from a bonebed that was inexpertly excavated and prepared, resulting in poorly preserved materials for study (Chure 1995). The specimen is similar to Allosaurus and has been referred to that genus (Paul 1988a; Smith 1998). However, Chure (1995) has identified at least two apomorphies on the skeleton. In addition, the locality appears to derive from high within the Morrison Formation, potentially above most other Allosaurus sites (Turner & Peterson 1999). We retain it here as a separate taxon. Shaochilong maortuensis (Hu, 1964) 1964 Chilantaisaurus maortuensis Hu: 50, figs 9–12, pls 1, 2. 2009a Shaochilong maortuensis (Hu); Brusatte et al.: 1052, figs 1, 2.

Holotype. LDM-LCA 9701-IV, a partial skull and skeleton. Diagnosis. Allosauroid with supraoccipital excluded from foramen magnum by midline contact between exoccipitals (otherwise unknown in tetanuran theropods; Wu et al. 2009). Occurrence. Quarry 1, Laochangjing, A’na village, Chuanjie township, Lufeng County, Yunnan, China; base of Upper Lufeng Formation; Middle Jurassic (Wu et al. 2009). Remarks. The original description of Shidaisaurus (Wu et al. 2009) listed a unique combination of several features, including downturned paraoccipital processes, the morphology of the lamina between the axial neural spine and epipophysis, proportions of the pelvic elements and

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absence of a notch distal to the ischial obturator process. However, these are present generally among sinraptorids and other tetanurans and are best dealt with as descriptive character states outside of the taxon diagnosis. In addition, although Wu et al. (2009, p. 9) stated that pleurocoels were not present in “any anterior dorsal vertebrae” of Shidaisaurus, they also noted that none of the vertebrae prior to dorsal 4 are exposed. Given that most theropods express pneumaticity up to but not past dorsal 4, the presence of this potential autapomorphy among noncoelurosaurian theropods cannot be determined. Siamotyrannus isanensis Buffetaut et al., 1996


1996 Siamotyrannus isanensis Buffetaut et al.: 689, figs 1, 2. Holotype. PW9-1, posterior dorsal vertebrae, sacrum, pelvis, and anterior caudal vertebrae (cast in Musée des Dinosaures, Esperaza). Diagnosis. Avetheropod with double vertical ridge on central part of lateral iliac blade (Rauhut 2003). Occurrence. Phu Wiang 9, Amphoe Phu Wiang, Changwat Khon Kaen, Thailand; Sao Khua Formation; Barremian–Aptian, Early Cretaceous (Racey & Goodall 2009). Remarks. In their original description, Buffetaut et al. (1996) interpreted Siamotyrannus as an Early Cretaceous tyrannosaurid, noting particularly the development of vertical ridging on the lateral surface of the ilium. However, this feature is expressed to varying degrees of prominence among widespread basal tetanurans other than tyrannosauroids, including Allosaurus, ‘Iliosuchus’, Megalosaurus and Piatnitzkysaurus (Bonaparte 1986; Benson 2009b, 2010a). The preserved elements of Siamotyrannus differ from corresponding tyrannosaur bones in several respects, especially the pubis and ischium (Fig. 6). For example, the ilium is broken distally in Siamotyrannus, not naturally abbreviated as in tyrannosaurs and other coelurosaurs. In fact this bone is quite robust even up to the preserved break surface. The ischium further bears a midshaft bulge developed by posterior extension of the contacting medial surfaces of the ischia, seen elsewhere in Sinraptor and Yangchuanosaurus. The pubic obturator foramen is open, but not broadly so as in tyrannosaurids, instead resembling the unusual condition exhibited by Sinraptor and Yangchuanosaurus. Likewise, the pubic boot is oriented similarly to that in Sinraptor (strongly posterodorsally inclined). With a thick anterior edge and a narrower posterior portion, it is much closer to the condition seen in basal tetanurans than coelurosaurs such as tyrannosauroids.

Figure 6. Left pelvic girdle of Siamotyrannus isanensis Buffetaut et al., 1996 (Musée des Dinosaures cast of PW9-1), showing features discussed in text. Abbreviations: ip, ischial peduncle; pb, pubic boot; pof, pubic obturator foramen; vr, vertical iliac ridges.

Although the ilium does show a pair of vertical ridges, these are much fainter in Siamotyrannus than in tyrannosauroids, where a single prominent ridge is typical. The anterior edge lacks the bilobate shape seen in most tyrannosaurids. Finally, although the pubic peduncle of the ilium is larger than the iliac peduncle, as is typical of tetanurans, the latter is not acuminate in Siamotyrannus, as would be typical of coelurosaurs. Sinraptor dongi Currie & Zhao, 1994 1994 Sinraptor dongi Currie & Zhao: 2039, figs 2–28. 1999 Yangchuanosaurus dongi (Currie & Zhao); Gao: 3. Holotype. IVPP 10600, complete skull and partial skeleton. Diagnosis. Allosauroid with: (1) enlarged lateral temporal fenestra with relatively straight postorbital-squamosal bar; (2) very short squamosal ramus of postorbital; and (3) palatine very deeply pneumatic between internal naris and postpalatine fenestra (all from Currie & Zhao 1994).



Occurrence. Jianjungmiao, Junggar Basin, Xinjiang Uygur Zizhiqu, China; upper part, Shishugou Formation; Oxfordian, Late Jurassic.

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Remarks. The well-preserved, complete skull and associated partial skeleton of Sinraptor have been described and illustrated in detail, providing a keystone for the recognition of an entire clade of theropods, Sinraptoridae (Currie & Zhao 1994). Although Sinraptor shares many features with allosauroids, it also retains a more primitive morphology in the manus and pelvis that resembles the condition in megalosauroids.

More recent discoveries across North Africa (Buffetaut 1989, 1992; Buffetaut & Ouaja 2002; Russell 1996; Dal Sasso et al. 2005) have brought to light new materials of Spinosaurus that, along with specimens of allied forms, have greatly illuminated its anatomy and relationships. Some materials have been assigned to a distinct genus, S. maroccanus (Russell 1996; Taquet & Russell 1998), but other workers have subsumed this within S. aegyptiacus (Sereno et al. 1998). We follow the latter course here, but note that, as with Carcharodontosaurus, future discoveries may better support the presence of distinct species in the differently aged North African beds.

Spinosaurus aegyptiacus Stromer, 1915

Streptospondylus altdorfensis von Meyer, 1832

1915 Spinosaurus aegyptiacus Stromer: 28, pls 1, 2. 1996 Spinosaurus maroccanus Russell: 355, figs 4–8. Holotype. BSP 1915, a partial skull and skeleton (destroyed). Hypodigm. Holotype; NHMUK R16420, the anterior portion of a skull, R16421, the anterior end of a dentary; MSNM V4047 (S. cf. S. aegyptiacus), a partial skull; MNHN SAM 124-128 (type, S. maroccanus), fragmentary jaws; and UCPC 2, a midline nasal crest. Diagnosis. Spinosaurid with: (1) no midline crest on conjoined premaxillae (Dal Sasso et al. 2005); (2) premaxilla entirely excluded from borders of external naris (Dal Sasso et al. 2005); and (3) extremely elongate dorsal neural spines (Stromer 1915). Occurrence. Baharije Oasis, Egypt and Kem Kem region, Morocco; Baharije Formation and Kem Kem beds; Albian?–early Cenomanian, Late Cretaceous. Remarks. Like Carcharodontosaurus, Spinosaurus is based on materials that survive only as lithographic plates (Stromer 1915), the originals having been destroyed in the same World War II raid on Munich. However, unlike Carcharodontosaurus, Spinosaurus did not enter into a period of obscurity thanks to the highly unusual nature of the type specimen. Characterized by enormously elongate neural spines, Spinosaurus and the family Spinosauridae have always retained a place in both scientific and popular literature. This lasting fame has helped perpetuate the highly speculative reconstruction made by Huene (1956, fig. 517) and the notion that Spinosaurus may have been partly quadrupedal (Charig & Milner 1986). With little anatomical data available, most researchers either placed it alone within its own family (e.g. Stromer 1915; Huene 1926b; Piveteau 1955) or allied it with the only other long spined genera then known, Acrocanthosaurus and Altispinax (e.g. Walker 1964; Romer 1966; Steel 1970).

1832 Streptospondylus altdorfensis von Meyer: 106. 1842 Streptospondylus cuvieri Owen: 88. 1867 Laelaps gallicus Cope: 234. 1908 Megalosaurus cuvieri (Owen); Huene: 332, figs 312, 313. 1964 Eustreptospondylus divesensis Walker: 124. Holotype. MNHN 8605–8609, 8787–8789, 8793, 8794, 8907, axial and appendicular elements from a single individual. Hypodigm. Holotype and MNHN 9645, the distal part of a left femur. Diagnosis. Tetanuran with bifurcated (or paired) hypapophyses on ventral surface of anterior dorsal vertebrae (Allain 2001; Benson 2010a, fig. 19B). Occurrence. Calvados, Falaises des Vaches Noires, near Honfleur, Basse-Normandie, France; Marnes de Dives; late Callovian–early Oxfordian, late Middle–early Late Jurassic. Remarks. The partial remains of Streptospondylus were some of the earliest dinosaur bones formally described, albeit intermixed with (and interpreted as) crocodylian remains at the time (Cuvier 1808; Allain 2001). The material is very incomplete but comes from a poorly known time interval for theropod evolution and thus has potential importance. Although the presence of bifurcated hypapophyses has been used to unite Streptospondylus and Eustreptospondylus as sister taxa (Allain 2001), it is not present in the latter form and is here considered an autapomorphy of Streptospondylus. Suchomimus tenerensis Sereno et al., 1998 1998 Suchomimus tenerensis Sereno et al.: 1298, figs 2, 3. 2002 Baryonyx tenerensis (Sereno et al.); Sues et al.: 545. Holotype. MNN GDF 500, partial postcranial skeleton.

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Hypodigm. Holotype and MNN GDF 501–508, 510, 511, cranial and postcranial materials. Diagnosis. Spinosaurid with: (1) elongate posterolateral premaxillary process that nearly excludes maxilla from external naris; (2) posterior dorsal, sacral and anterior caudal vertebral neural spines expanded anteroposteriorly and dorsoventrally; and (3) hook-shaped radial ectepicondyle (all from Sereno et al. 1998).


Occurrence. Gadoufaoua outcrop, Agadez, Niger; Elrhaz Formation, Tegama Group; Aptian–Albian?, Early Cretaceous. Remarks. Suchomimus is one of the better represented spinosaurids, with well-preserved material known from the skull, axial skeleton, and appendicular region of several individuals over a range of body sizes. There has been discussion regarding the validity of Suchomimus as a distinct genus from Baryonyx, with some (Sues et al. 2002) favouring referral of S. tenerensis to the latter genus. Among the diagnostic features listed above for Suchomimus, character (1) cannot be observed in Baryonyx and (2) is difficult to assess because of the immature nature of the Baryonyx holotype. (A later study suggested that the neural spines over the pelvic region may also be expanded in Baryonyx [Charig & Milner 1990].) It remains subjective whether enlargement of the radial entepicondyle justifies generic separation, but we retain the original designation here. A fourth autapomorphy, presence of a hypertrophied ulnar olecranon process offset from the humeral articulation, was also listed originally (Sereno et al. 1998) but appears to be present in Baryonyx, although obscured by incomplete preservation of the proximal ulna. ‘Szechuanosaurus’ zigongensis Gao, 1993 1993 Szechuanosaurus zigongensis Gao: 308, figs 1–6, pls 1–3. Holotype. ZDM 9011, a partial postcranial skeleton. Hypodigm. Holotype, ZDM 9012, a left maxilla; 9013, two teeth; and 9014, a right hind limb (Gao 1993). Diagnosis. Tetanuran with only cervical vertebrae 2–4 opisthocoelous, remainder amphiplatyan (Holtz et al. 2004). Although the specimen requires detailed restudy, it can be distinguished from other theropods in the same formation (Rauhut 2003). Occurrence. Dashanpu Dinosaur Quarry, Zigong, Sichuan, China; Xiashaximiao Formation; Middle Jurassic (Gao 1993).

Remarks. The holotype skeleton of ‘S.’ zigongensis has been included in a number of phylogenetic studies (Rauhut 2003; Holtz et al. 2004), which have tended to recover it among basal tetanurans. Unfortunately, the holotype of Szechuanosaurus campi, the type species of the genus, is non-diagnostic and therefore S. campi is a nomen dubium (Chure 2001a; Holtz et al. 2004). The species ‘S.’ zigongensis, however, is based on much of a postcranial skeleton that Chure (2001a) felt was sufficient to justify creation of a new taxon to include it and CV 00214. We do not consider these forms close enough to include in one taxon, and indeed they derive from different strata and can be scored differently for many of the characters in our analysis. We therefore retain ‘S.’ zigongensis as a distinct OTU. Torvosaurus tanneri Galton & Jensen, 1979 1979 Torvosaurus tanneri Galton & Jensen: 1, figs 1, 2, 3A, G, L, 5–7, 8H. 1988a Megalosaurus tanneri (Galton & Jensen); Paul: 282. 1992 Edmarka rex Bakker et al.: 2, figs 1, 3, 7, 10, 12–15. 1997 ‘Brontoraptor’ sp. Siegwarth et al.: 4, figs 1–9, 10A–E, 11A–E, 12–13A, 14–15A, 16A–H, 17 (nomen nudum). Holotype. BYU-VP 2002, left and right forelimbs. Hypodigm. Holotype and BYU-VP 2000 series; TATE 401, 1002–1005 (holotype, Edmarka rex), jugal, scapulocoracoid, and ribs (Bakker et al. 1992); 1003, 1012 (‘Brontoraptor’), pelvis, femur? and other postcranial material (Siegwarth et al. 1997). Diagnosis. Megalosauroid with: (1) very shallow maxillary fossa; (2) ‘fused’ paradental plates (Britt 1991); (3) pronounced rim around anterior face of cervical centra (Britt 1991); (4) expanded fossae in posterior dorsal and anterior caudal vertebrae centra forming enlarged, deep openings (modified from Britt 1991) that we interpret as pneumatic; (5) highly ossified puboischiadic plate (modified from Britt 1991); and (6) distal expansion of ischium with prominent lateral midline crest and oval outline in lateral view. Occurrence. Dry Mesa Quarry, Montrose County; Calico Gulch Quarry, Uncompahgre Plateau, Moffit County, and Meyer site, Garden Park, north of Cañon City, Fremont County, Colorado; Carnegie Quarry, Dinosaur National Monument, Uintah County, Utah; Gilmore Quarry N, Freezeout Hills, Carbon County, and Nail and Louise Quarries, Como Bluff, Albany County, Wyoming, USA; Salt Wash and Brushy Basin Members, Morrison Formation; Kimmeridgian–Tithonian, Late Jurassic. Remarks. Torvosaurus was an important discovery from the Morrison Formation (Galton & Jensen 1979), not only


The phylogeny of Tetanurae because it represented a new large theropod, but also because it appeared to be more primitive than Allosaurus and in many ways similar to Megalosaurus (Galton & Jensen 1979; Britt 1991). Later, additional specimens were discovered at other Morrison Formation quarries. Among these, the incomplete materials that have been referred to Edmarka rex (Bakker et al. 1992) and ‘Brontoraptor’ (Siegwarth et al. 1997) may represent species level variants of Torvosaurus, but we do not consider the observed differences as sufficient to justify a new taxon at this time. Therefore we consider E. rex and ‘Brontoraptor’ as junior synonyms of T. tanneri (Holtz et al. 2004). Mateus and Antunes (2000) referred a tibia (ML 430) from the early Tithonian of Casal do Bicho, Portugal to Torvosaurus sp. It is likely that this does indeed represent Torvosaurus, although it may also belong to a related form; there are no clear autapomorphies on the tibia aside from its very robust proportions. Mateus et al. (2006) later referred a maxilla (ML 1100) from the base of the Lourinhã Formation of Praia da Vermelha, Portugal to Torvosaurus tanneri. However, they noted a much reduced tooth count of 10 (compared to 11–13 in the topotype maxilla), which suggests a more conservative assignment to Torvosaurus sp. pending the discovery of further material. Tyrannotitan chubutensis Novas et al., 2005 2005 Tyrannotitan chubutensis Novas et al.: 226, figs 1, 2. Holotype. MPEF-PV 1156, a partial skeleton. Hypodigm. Holotype and MPEF-PV 1157, a partial skeleton. Diagnosis. Allosauroid with: (1) bilobate denticles on anterior carinae of teeth; and (2) very pronounced mental groove on lateral surface of dentary (all from Novas et al. 2005). Occurrence. Estancia ‘La Juanita’, 28 km north-east of Paso de Indios, Chubut, Argentina; Cerro Castaño Member, Cerro Barcino Formation; Aptian–Albian, Early Cretaceous (Rauhut pers. comm. 2011). Remarks. Slightly older than Giganotosaurus and comparable in size, Tyrannotitan is among the more recently described carcharodontosaurids (Novas et al. 2005). Although the holotype and referred specimen do not include the entirety of the skeleton, the preserved portions of the skull, axial, and appendicular regions show a number of carcharodontosaurid features as well as strong similarities with Giganotosaurus and Mapusaurus. MPEF-PV 1156 also provides evidence of forelimb shortening in derived carcharodontosaurids (Novas et al. 2005).

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Xuanhanosaurus qilixiaensis Dong, 1984 1984 Xuanhanosaurus qilixiaensis Dong: 213, figs 1–4, pl. 1. Holotype. IVPP V.6729, partial forelimbs and anterior and middle dorsal vertebrae. Diagnosis. Tetanuran with: (1) pronounced posterior ridge on articular facet of humeral head that overhangs shaft (Rauhut 2003); and (2) dorsal neural spines transversely thick with gently concave lateral surfaces (Rauhut pers. comm. 2007). Occurrence. Xuanhan County, Sichuan, China; Xiashaximiao Formation; Middle Jurassic. Remarks. Known only from an associated forelimb and vertebrae, Xuanhanosaurus nevertheless appears to preserve enough morphological information both to distinguish it from other theropods and to inform its phylogenetic position. Previously it has been recovered as a basal tetanuran (Holtz et al. 2004) and a megalosauroid (Benson 2010a). The robust humerus and wide scapular blade are accompanied by a four-digit manus with relatively short metacarpals. This unusual combination of features suggests a basal position within Tetanurae, and indeed the manus bears some resemblance to those of ‘Szechuanosaurus’ zigongensis and Torvosaurus. ‘Yangchuanosaurus’ hepingensis Gao, 1992 1992 Yangchuanosaurus hepingensis Gao: 313, figs 1–4, pls 1–3. 1994 Sinraptor hepingensis (Gao); Currie & Zhao: 2039. Holotype. ZDM T0024, complete skull and skeleton. Diagnosis. As with Yangchuanosaurus shangyouensis (q.v.), it is not clear whether ‘Yangchuanosaurus’ hepingensis possesses autapomorphies or can only be distinguished by a unique combination of characters (see Remarks, below). Occurrence. Dashanpu Dinosaur Quarry, Zigong, Sichuan, China; Shangshaximiao Formation; Oxfordian– early Kimmeridgian, Late Jurassic. Remarks. The ‘sinraptorid’ species hepingensis has alternately been referred to the genera Yangchuanosaurus (Gao 1992, 1999) and Sinraptor (Currie & Zhao 1994; Rauhut 2003; Holtz et al. 2004), depending on the perceived importance of particular character states. For example, the proportionally long, low skull and very tall dorsal vertebral neural spines are more similar to Sinraptor (Currie & Zhao 1994), whereas the sinuous, rugose nasal crest, marked margin of

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the jugal antorbital fossa, and more horizontally oriented pubic boot resemble the conditions in Yangchuanosaurus. Given this apparent mosaic of character states, we retain ‘Y .’ hepingensis as a discrete taxon in our analysis. Yangchuanosaurus shangyouensis Dong et al., 1978 1978 Yangchuanosaurus shangyouensis Dong et al.: 302, figs 1, 2. 1983 Yangchuanosaurus magnus Dong et al.: 83, pls 27–31, figs 54–63. 1988a Metriacanthosaurus shangyouensis (Dong et al.); Paul: 290.


Holotype. CV 00215, a complete skull and skeleton. Hypodigm. Holotype and CV 00216, a complete skull and skeleton (holotype, Y. magnus). Diagnosis. The original diagnoses of Yangchuanosaurus and its two constituent species (Dong et al. 1978, 1983) include features that now characterize ‘sinraptorids’, allosauroids and tetanurans more generally. More recent work (Currie & Zhao 1994) has distinguished Yangchuanosaurus from Sinraptor primarily on the presence of sinraptorid characters together with the absence of Sinraptor autapomorphies (i.e. a unique character combination but no autapomorphies). Yangchuanosaurus has a slightly higher ratio of skull height to length (0.5) than Sinraptor (0.4) and correspondingly has a proportionally taller maxilla. The dorsal vertebral neural spines are also lower (about 1.8 times centrum height compared to 2.0 in S. dongi) and the centra are relatively longer. Yangchuanosaurus may also exhibit a more pronounced margin of the antorbital fossa on the jugal, although this is difficult to ascertain in some sinraptorid specimens. We concur that Yangchuanosaurus can be distinguished from other sinraptorids, but in the absence of a detailed restudy of the type materials we can offer no new autapomorphies to define this genus. Occurrence. Shangyou Reservoir and Hongjiang Machine Factory, near Yongchuan, Yongchuan County, Sichuan, China; Shangshaximiao Formation; Oxfordian–early Kimmeridgian, Late Jurassic. Remarks. The original and emended diagnoses of Yangchuanosaurus shangyouensis and Y. magnus differentiated these species primarily on the basis of size (Dong et al. 1978, 1983). In addition, Dong et al. (1983) noted that the maxilla of Y. magnus housed an additional fenestra within the antorbital fossa, whereas Y. shangyouensis possessed only a fossa in this location. Given the tendencies towards increased pneumatization in larger (older) individuals among theropods (Wedel 2003; although see Rauhut & Fechner 2005; Brusatte et al. 2009b) and the variability of pneumatic features even within a single individual (Zhao &

Currie 1994), we consider this likely to be an intraspecific, possibly ontogenetic, variation. The apparent difference in cervical vertebral morphology (ventral keels absent in Y. shangyouensis, present in Y. magnus) may be a function of comparing different positions within the column. Therefore, the holotypes of the two species of Yangchuanosaurus are effectively identical, and indeed they would have the same patterns of character codings in our matrix. Given their provenance and the high degree of similarity between these specimens, we consider them to represent a single species.

Phylogenetic analysis Characters This study employed a total of 351 characters derived from a combination of previous phylogenetic studies (see online Supplementary Table 1) and direct study of specimens by the authors. Numerous previously described characters were not employed in this study, and it would be a gargantuan task to discuss these individually. In general, ‘excluded’ characters were: (1) parsimony uninformative for our taxon set; (2) redundant or subsumed into existing characters; or (3) not explicable or observable by the present authors. All included characters were scored based on direct examination of fossil materials or casts, except in those cases where key specimens no longer exist or where we could not visit for study. In total, more than 90% of included taxa were examined directly by one or more of the authors. Character descriptions are listed in Appendix 3 (see Supplementary Material online). The taxon-character matrix is given in Appendix 2, and executable version is available online as Supplementary File 3.

Phylogenetic methods The 351 characters used in this analysis were unordered and equally weighted, and scored using Mesquite 2.72 (Maddison & Maddison 2009). The resulting matrix was imported into TNT 1.1 (Goloboff et al. 2008). Given the large size of this matrix, we opted for a heuristic search using the ‘New Technology’ options. These included the default settings for sectorial, ratchet, tree drift and tree fusion, using a driven search that stabilized consensus twice with a factor of 25. Subsequently, we subjected the resulting most parsimonious trees (MPTs) to tree bisection and reconnection (TBR) branch swapping. The trees resulting from both search iterations were examined using strict and Adams consensus (‘combinable components’ in TNT). We confirmed the results of this analysis using PAUP∗ 4.0b10 (Swofford 2002), employing the parsimony ratchet (Nixon 1999) implemented in PAUPRat (Sikes & Lewis 2001) and TBR branch swapping on the resulting trees. We isolated ‘wildcard’ taxa using the Adams consensus tree. ‘Wildcards’ are taxa that take multiple phylogenetic



placements with equal parsimony, thus resulting in a poorly resolved strict consensus. Pruning wildcards from the set of MPTs and then calculating the strict consensus results in a ‘strict reduced consensus’. This strategy summarizes relationships among non-wildcard taxa that are otherwise concealed by inclusion of wildcards (Wilkinson 2003). We performed successive iterations of our search strategy on the same character matrix after having removed these taxa. Following the initial analyses, we determined branch support using the ‘Decay Index PAUP File’ function of MacClade (Maddison & Maddison 1992) combined with the parsimony ratchet implemented using PAUPRat in PAUP∗ to determine the shortest tree length lacking each clade that was originally recovered by our analysis.

Stratigraphic fit methods We employed three commonly used methods to assess the stratigraphic fit of our phylogenetic results. We used a Spearman rank correlation to test the relationship between age rank (using stages, numbered from oldest to youngest) and clade rank (numbering nodes upwards from the base of the tree), although we did not collapse the cladogram into a ‘comb’ topology. The Stratigraphic Consistency Index (SCI; Huelsenbeck 1994) was used to assess the number of nodes that were consistent with the stratigraphic placement of the oldest included taxa. Finally the Relative Completeness Index (RCI; Hitchin & Benton 1997) measures the amount of record missing based on the current phylogeny.

Results Tree length and topology The initial TNT analysis produced 91 most parsimonious trees, each of 1020 steps; subsequent TBR branch swapping resulted in fewer than 40,000 trees of 1020 steps each. The consistency index (CI) was 0.4216 (HI therefore = 0.5784), the retention index (RI) = 0.6956, and the rescaled consistency index (RC) = 0.2932. Excluding parsimony uninformative characters, the CI was 0.2404 and the HI was 0.5796. Results were identical when PAUP∗ was used to perform the search, but more than 400,000 MPTs were recovered. A strict consensus of these trees produced the cladogram in Fig. 7A. This phylogeny provides significant resolution among the major tetanuran clades that confirms the successive placement of coelophysoids, ceratosaurs and tetanurans within Theropoda. Dilophosaurus is recovered as a coelophysoid, in contrast to a recent study (Nesbitt et al. 2009), whereas Cryolophosaurus and ‘D.’ sinensis are more derived. Tetanurae is monophyletic and includes most of its traditional subclades. Within Tetanurae we recovered: (1) a basal clade, Megalosauroidea (= Spinosauroidea, see below), which includes

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spinosaurids, megalosaurids and a third, more basal clade; (2) a more derived Allosauroidea, including allosaurids, sinraptorids (= metriacanthosaurids, see below), neovenatorids and carcharodontosaurids; and (3) Coelurosauria, here represented by Compsognathus, Ornitholestes and Proceratosaurus, the sister taxon to Allosauroidea. Megalosauroidea is composed of three main clades. Among them, Spinosauridae includes Spinosaurinae (Spinosaurus, Irritator and Angaturama) and Baryonychinae (Baryonyx and Suchomimus). Its sister lineage Megalosauridae is a diverse assemblage that includes at least two clades, Megalosaurus + Torvosaurus + Duriavenator and a poorly resolved second group (Dubreuillosaurus, Afrovenator, Piveteausaurus, Leshansaurus, Magnosaurus and Poekilopleuron); Eustreptospondylus and Streptospondylus are unresolved within Megalosauridae + Spinosauridae. A Piatnitzkysaurus + Marshosaurus + Condorraptor clade is found to be basal within Megalosauroidea. Among the allosauroids, ‘sinraptorids’ are the most basal group. The clade is more diverse than has previously been assessed, with six taxa aggregated into two groups (Metriacanthosaurus + Sinraptor dongi + Siamotyrannus + ‘Y.’ hepingensis and Yangchuanosaurus shangyouensis + ‘S.’ zigongensis + CV 00214) plus two unresolved basal forms (Shidaisaurus and Xuanhanosaurus). Allosauridae exists only as Allosaurus + Saurophaganax, the sister lineage to Carcharodontosauria (Carcharodontosauridae + Neovenatoridae). Several recently discovered forms make this latter the most diverse allosauroid clade. Neovenatoridae comprises Neovenator, Chilantaisaurus, Aerosteon + Megaraptor, and Australovenator + Fukuiraptor. The remaining taxa (Eocarcharia, Concavenator, Shaochilong, Acrocanthosaurus, Mapusaurus, Tyrannotitan, Carcharodontosaurus and Giganotosaurus) form Carcharodontosauridae. This analysis recovered several individual taxa in wellresolved positions outside major clades or ingroups. Among them, Cryolophosaurus and ‘D.’ sinensis are basally placed ‘stem’ tetanurans; Monolophosaurus and Chuandongocoelurus occupy similar but slightly more derived positions. Interestingly, Lourinhanosaurus is allied with Coelurosauria. An Adams consensus (Fig. 7B) identified Poekilopleuron, Streptospondylus and Xuanhanosaurus as ‘wildcard’ taxa. Deletion of Streptospondylus resulted in a fourfold reduction to 105,750 unique trees and the resolution of Eustreptospondylus as the most basal megalosaurid. Subsequent deletion of Xuanhanosaurus resulted in an additional five-fold reduction to 21,150 unique trees and resolution of Shidaisaurus as a basal representative of Metriacanthosaurinae; eliminating Poekilopleuron resulted in 4050 trees, with no further improvement in resolution (Fig. 8A). Finally, we conducted an Adams consensus on the 4050 trees resulting from the pruned analysis. This topology



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Figure 7. Results of current phylogenetic analysis. A, strict consensus result. Names in bold refer to newly discovered or significantly redefined clades resulting from this analysis. Numbers to the left of the nodes indicate unambiguous character support, those to the right show branch support. B, Adams consensus result. Names and lines in bold indicate potential ‘wildcard’ taxa’.

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Figure 8. Results of phylogenetic analysis after pruning Poekilopleuron, Streptospondylus and Xuanhanosaurus from the full set of most parsimonious trees. Numbers at nodes indicate branch support. A, strict consensus result. B, Adams consensus result.

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(Fig. 8B) suggests that several additional taxa may be acting as ‘wildcards’, but their removal does not improve resolution beyond that achieved by deleting one taxon from any polytomy.


Branch support and taxon instability Character support for each node is reported in online Supplementary File 3. Branch support is low (≤2) for many nodes, but several clades (Coelophysidae, Neotheropoda, Averostra, Ceratosauria, Spinosauridae) show significant support (> 5). However, in many instances the low support is due to the uncertain position of a single taxon. These are typically fragmentary or otherwise poorly known forms whose instability gives a spuriously low confidence value for the overall scheme of relationships. For example, a single added step allows the following placements: Chilantaisaurus as a basal carcharodontosaurid close to Concavenator; Shidaisaurus as a basal metriacanthosaurid or the sister taxon to Metriacanthosaurus; Monolophosaurus as a basal megalosauroid; and Cryolophosaurus outside Averostra or just above ‘D.’ sinensis. Lourinhanosaurus can be recovered as a basal allosauroid or a basal avetheropod with the addition of two steps. In addition, we recover Poekilopleuron as an afrovenatorine megalosaurid, but with a single additional step it can be recovered as a piatnitzkysaurid, elsewhere within Megalosauridae, or at several possible positions within Allosauria. This effectively reduces support for the entire avetheropod ‘stem’, many allosaurian ingroups, and much of Megalosauroidea down to 1. A significant improvement in overall support is actually achieved if Poekilopleuron is removed from the analysis and branch support is recalculated. In this case, support for several nodes rises significantly, including Allosauria and Allosauroidea (to 3), and Metriacanthosauridae and Carcharodontosauria (to 4), while most other nodes now have a support of at least 2. Given how fragmentary the only known specimen of Poekilopleuron is, we consider its placement within Afrovenatorinae to be extremely tentative, and realistically it could represent another megalosaurian or even an allosaurian.

Taxonomic and nomenclatural implications The topologies produced by our analyses indicate several changes to existing taxonomic usage. These are described below, and summarized in Appendix 1. The family Megalosauridae was established in Fitzinger (1843; non Huxley 1870) in accordance with ICZN Article 11.7.2. It takes priority over Spinosauridae Stromer, 1915 and Torvosauridae Galton & Jensen, 1979; therefore Megalosauroidea is a senior synonym of Spinosauroidea and Torvosauroidea. It is used here to include all taxa closer to Megalosaurus than to Allosaurus or Tyrannosaurus. Within this clade, Spinosauridae and Megalosauridae are

most closely related; we append the existing name Megalosauria Bonaparte, 1850 to this node. Megalosaurinae can be defined as all megalosaurids closer to Megalosaurus than to Afrovenator. Metriacanthosaurus is the type genus of Metriacanthosauridae Paul, 1988a, which was defined only briefly but with purportedly differential characters (in accordance with ICZN Article 13.1.1). The robust recovery of Metriacanthosaurus within Sinraptoridae Currie & Zhao, 1994 places the latter family name into synonymy with the former. Two lineages are evident within Metriacanthosauridae, but individual taxa are not interrelated in a manner that supports all previous species assignments. For example, we recover a sister-taxon relationship between Yangchuanosaurus shangyouensis and specimen CV 00214, with the type of ‘Szechuanosaurus’ zigongensis as outgroup to this pair. This contradicts the suggestion that the latter two specimens pertain to a single species (Chure 2001a). Rather, we assign CV 00214 to Y. shangyouensis given its close similarity and derivation from the same stratum and geographic region. Thus we agree with Paul (1988a) that it is congeneric with Yangchuanosaurus shangyouensis but do not agree that the differences warrant specific separation. We acknowledge the close relationship between ‘Szechuanosaurus’ zigongensis and Yangchuanosaurus by referring the former to Yangchuanosaurus as Y. zigongensis but retain it as a distinct species given its earlier provenance. The name Yangchuanosaurus can be applied to this entire clade. However, ‘Y.’ hepingensis is more closely related to Sinraptor than to Yangchuanosaurus, supporting its recent referral to the former genus (Currie & Zhao 1994). S. hepingensis and S. dongi belong to a lineage that also includes Metriacanthosaurus and Siamotyrannus, and can be defined as follows. Tetanurae Gauthier, 1986 Allosauroidea Currie & Zhao, 1994 Metriacanthosauridae Paul, 1988a Metriacanthosaurinae (Paul 1988a) subfam. nov. Type genus. Metriacanthosaurus Walker, 1964. Included taxa. Metriacanthosaurus parkeri (Huene, 1923) Walker, 1964; Siamotyrannus isanensis Buffetaut et al., 1996; Sinraptor hepingensis (Gao, 1992) Currie & Zhao, 1994; Sinraptor dongi Currie & Zhao, 1994. Shidaisaurus jinae Wu et al., 2009 might also belong here, based on the results of our reduced dataset analysis. Diagnosis. Metriacanthosaurids possessing the following synapomorphies: (1) anteroventral border of maxillary antorbital fossa demarcated by raised ridge (22(1); also in Eoraptor, Coelophysidae, Masiakasaurus, Marshosaurus and Compsognathus); (2) pronounced ventral keel on



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anterior dorsal vertebrae (184(1)); also in Condorraptor, Piatnitzkysaurus, Carcharodontosaurus and many megalosaurians); (3) straight posterior margin of iliac postacetabular process (280(2)); (3) angle of less than 60◦ between long axes of pubic shaft and boot (285(1)); (4) ventrally curved ischial shaft (293(1)); also in Coelophysidae, Eustreptospondylus, Afrovenator, Megalosaurus and Compsognathus); and (5) bulbous fibular crest on tibia (323(1)); also in ‘Dilophosaurus’ sinensis).

Definition. All megalosauroids more closely related to Piatnitzkysaurus than to either Spinosaurus or Megalosaurus. Finally, we have recovered a novel clade within Megalosauridae that includes six taxa: Piveteausaurus, Leshansaurus, Afrovenator, Dubreuillosaurus, Poekilopleuron and Magnosaurus. No existing name can be applied to this clade, so we supply the following.

Definition. All metriacanthosaurids more closely related to Metriacanthosaurus than to Yangchuanosaurus.

Tetanurae Gauthier, 1986 Megalosauroidea Fitzinger, 1843 Megalosauridae Fitzinger, 1843 Afrovenatorinae subfam. nov.

Allosauridae is reduced to Allosaurus + Saurophaganax but holds a stable place as the sister taxon to Carcharodontosauria. We suggest using the existing name Allosauria Paul, 1988a for this node, in parallel with the nomenclatural arrangement outlined above for megalosauroids. The position of ‘D.’ sinensis as distinct from D. wetherilli is likely secure, as the two taxa are well separated in our results, and therefore this species requires assignment to a different genus. Forthcoming research (P. Currie pers. comm. 2009) will address this issue in more detail. We also recover a clade consisting of Piatnitzkysaurus, Marshosaurus and Condorraptor at the base of Megalosauroidea. These taxa show numerous morphological similarities as well as distinctions from other megalosauroids, and we consider it useful to provide a name for this grouping. Tetanurae Gauthier, 1986 Megalosauroidea (Fitzinger, 1843) Piatnitzkysauridae fam. nov. Type genus. Piatnitzkysaurus Bonaparte, 1979. Included taxa. Piatnitzkysaurus floresi Bonaparte, 1979; Marshosaurus bicentesimus Madsen, 1976b; Condorraptor currumili Rauhut, 2005a. Diagnosis. Megalosauroids possessing the following synapomorphies: (1) two parallel rows of nutrient foramina on lateral surface of maxilla (21(1)); also in Eocarcharia and Shaochilong); (2) vertically striated or ridged paradental plates (140(1)); also in abelisaurids, Megalosaurus and Proceratosaurus); (3) reduced axial parapophyses (166(1)); also in coelophysoids, Eustreptospondylus and Afrovenator); (4) anteriorly inclined posterior dorsal neural spines (192(1)); in parallel with Allosauroidea); and (5) canted distal humeral condyles (236(1)); also in Poekilopleuron, Allosauridae and Fukuiraptor). In addition, among tetanurans these taxa show reversals to the primitive condition for the following features: (1) a short or absent anterior maxillary ramus (12(0)); (2) moderate development of axial diapophyses (167(0)); and (3) no axial pleurocoels (168(0)).

Type genus. Afrovenator Sereno et al., 1994. Included taxa. Afrovenator abakensis Sereno et al., 1994; Dubreuillosaurus valesdunensis (Allain, 2002) Allain, 2005a; Leshansaurus qianweiensis Li et al., 2009; Magnosaurus nethercombensis (Huene, 1923) Huene, 1932; Piveteausaurus divesensis Walker, 1964; and tentatively Poekilopleuron bucklandii Eudes-Deslongchamps, 1837. Diagnosis. Megalosauroids possessing the following synapomorphies: (1) squared anterior margin of maxillary antorbital fenestra (23(1)); also in coelophysoids, Irritator, Concavenator and Eocarcharia); and (2) puboischiadic plate broadly open along midline (281(2)); in parallel with Avetheropoda). Definition. All megalosaurids more closely related to Afrovenator than to Megalosaurus. Finally, we establish the new name Orionides for the node comprising Megalosauroidea, Avetheropoda, their most recent common ancestor, and all its descendants. The name alludes to Orion, the giant hunter of Greek mythology (and to the alternate name for the constellation Orion, Alektropodion, or ‘chicken foot’).

Fragmentary taxa As with our analysis of ceratosaurs (Carrano & Sampson 2008), we did not include every potential basal tetanuran in this study; several were excluded because they were too fragmentary or provided no unique character combinations. The phylogenetic determinations discussed below are made to the most exclusive level possible. Although many of these forms cannot be identified beyond a rather general level (i.e. Allosauroidea), they nonetheless add important temporal and geographic data to the story of tetanuran evolution (Fig. 9). In keeping with the scope of this study, we have excluded coelurosaurs from the discussion below, and do not discuss every report of a theropod tooth unless it has been previously assigned to a specific basal tetanuran taxon.

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M. T. Carrano et al. Jurassic Early

He

Si

Pl

Cretaceous Middle

To

Aa

Bj

Bt Cl

Late Ox

Ki

Early Ti

Be Va

Ba

Late Ap

Al

Ce

Tu Co Sa

Ca

Ma

NORTH AMERICA

Saurophaganax

Ha

Acrocanthosaurus Arundel Cloverly

Allosaurus, Torvosaurus Marshosaurus

‘M.’ inexpectatus Cerro Lisando Alcântara

Aerosteon

Angaturama, Irritator

Condorraptor, Piatnitzkysaurus

Adamantina Unquillosaurus Afrovenator

Spinosaurus

Cabao

Allen

EUROPE

Eocarcharia Carcharodontosaurus ‘M.’ ingens Metriacanthosaurus Wadi Milk Mugher Mudstone Magnosaurus Kimmeridge Clay Baryonyx Duriavenator Allosaurus S. girardi Torvosaurus Piveteausaurus, Streptospondylus Neovenator, Baryonychinae Eustreptospondylus Enciso Cruxicheiros, Iliosuchus, Megalosaurus Hastings Morella Wessex Dubreuillosaurus, Poekilopleuron Pinilla de los Moros, Blesa Villar del Arzobispo Jet Rock Erectopus Montmirat ‘S.’ cuvieri Artoles Valdoraptor Lower Lias Becklespinax ‘Saltriosaurus’

‘M.’ pannoniensis

Fukuiraptor

Gasosaurus, Kaijiangosaurus, Shidaisaurus, Xuanhanosaurus, Y. zigongensis

ASIA

Chilantaisaurus, Shaochilong Leshansaurus, S. hepingensis Khok Kruat Y. shangyouensis S.dongi Siamosaurus, Siamotyrannus

Phu Kradung Chuandongocoelurus, Monolophosaurus Balabansai

‘A.’ sibiricus, Embasaurus

Jobu

Kelmayisaurus

Cryolophosaurus

Hidden Lake

Rapator Eumeralla

Australovenator

ANTARCT/ AUST

‘D.’ sinensis

AFRICA

Cristatusaurus, Suchomimus

Kirkwood

‘A.’ tendagurensis, Carcharodontosauria


Orkoraptor Marília

SOUTH AMERICA

Giganotosaurus Mapusaurus Megaraptor

Tyrannotitan

Figure 9. Stratigraphical ranges of non-coelurosaurian tetanurans. Symbols: open circles, basal or indeterminate theropods; filled circles, basal or indeterminate tetanurans; open triangles, piatnitzkysaurids, megalosaurids and indeterminate megalosauroids; filled triangles, spinosaurids; open squares, metriacanthosaurids, allosaurids and indeterminate allosauroids; filled squares, carcharodontosaurians.

A complete listing is presented in Supplementary Table 4 (see online Supplementary File 2). Allosaurus? sibiricus Riabinin, 1915. Described as the distal end of a right metatarsal IV from the Neocomian (Early Cretaceous) Turgin (?) Formation of Tarbagatai, Transbaikalia, Chitinskaya Oblast, Russia (Riabinin 1915; Nessov 1995), this bone is actually a left metatarsal II. It is too fragmentary to be assigned to a known taxon or identified as a distinct form, especially considering that this bone is not particularly diagnostic among non-coelurosaurian theropod clades. Its morphology is quite similar to that of Allosaurus and Neovenator but also to more primitive forms such as Afrovenator and Torvosaurus. Holtz et al. (2004) referred to it as Chilantaisaurus sibiricus, but it is best identified as Theropoda indet. Allosaurus? tendagurensis Janensch, 1925. A single tibia from the late Kimmeridgian–Tithonian Middle Dinosaur

Member of Tendaguru, Tanzania (MB.R 3620 = tibia 67) was made the holotype of this species by Janensch (1925). Although incomplete, and probably restored as too elongate, the proximal and distal ends are sufficiently well preserved. As suggested by Rauhut (2005b), this specimen clearly pertains to a tetanuran; it lacks the distinctive ceratosaur cnemial crest, the distal end bears a marked buttress for the (presumably laminar) astragalar ascending process, and the distal profile is transversely elongate. However, the specimen is not particularly similar to that of Allosaurus among tetanurans and cannot be more specifically identified. The assignment of isolated caudal vertebrae from Quarry TL to this taxon (Janensch 1925) is not justified. Becklespinax altispinax (Paul, 1988a) Olshevsky, 1991. Three dorsal vertebrae (NHMUK R1828; Fig. 10) from the Hastings Beds Group (late Berriasian–Valanginian) of Battle, England constitute the controversial type specimen


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showing only weak lateral indentations that are widely present among amniotes, and the lateral surfaces of the neural spines do not bear deep longitudinal troughs (derived carcharodontosaurids only). The spines are unusually thick, approximately as broad mediolaterally as they are long anteroposteriorly, and widen considerably toward their apices. They also show evidence of a healed break as well as some ossification of the interspinous (presumably ligamentous) soft tissues. The centrum proportions are similar to those of many non-coelurosaurian tetanurans. Other than to exclude Becklespinax from Spinosauridae and Carcharodontosauridae, we cannot place it more specifically within Tetanurae. Its exclusion from these two clades implies the independent derivation of elongate dorsal neural spines at least three times within Theropoda. Cristatusaurus lapparenti Taquet & Russell, 1998. Cristatusaurus lapparenti was the first spinosaurid described from the same formation (Elrhaz Formation; Aptian–Albian) and general location as Suchomimus (Taquet 1976; Taquet & Russell 1998). It has been widely considered a nomen dubium (Sereno et al. 1998; Sues et al. 2002; but see Allain 2002) because the holotype material (MNHN GDF 366) is extremely fragmentary and the accompanying diagnosis very general, a conclusion with which we agree. Cristatusaurus can only be identified as an indeterminate baryonychine spinosaurid. However, we also consider it unlikely that two baryonychines are present at Gadoufaoua, and therefore Cristatusaurus and Suchomimus almost certainly represent the same animal. Figure 10. Posterior dorsal vertebrae (NHMUK R1828) of Becklespinax altispinax (Paul, 1988a) in left lateral view. Abbreviations: hs, hyposphene; pb, pathological bone; poz, postzygapophysis; prz, prezygapophysis. Scale bar = 10 cm.

of this taxon. Originally, these materials were conditionally assigned to Megalosaurus dunkeri Dames, 1884 under the new genus Altispinax Huene, 1923. Specifically, Huene (1926a, p. 78) remarked that if these vertebrae belonged to the same animal as the holotype tooth of M. dunkeri, then the resulting taxon could be called Altispinax dunkeri. As this tooth is non-diagnostic within Theropoda (see below), and in any case cannot be assigned to the same taxon as the vertebrae, NHMUK R1828 was made the holotype of a new taxon (Paul 1988a; Olshevsky 1991). The vertebrae are unusual in possessing highly elongate neural spines, which has led many workers (Walker 1964; Romer 1966; Carroll 1988; Kurzanov 1989) to assign Becklespinax to Spinosauridae. However, they show important differences with the vertebrae of both spinosaurids and the long-spined carcharodontosaurids. Unlike spinosaurids, the arches lack accessory laminae either below or above the transverse process. Unlike carcharodontosaurids (and Torvosaurus), the centra are poorly or non-pneumatized,

Cruxicheiros newmanorum Benson & Radley, 2010. A recently described fragmentary skeleton from the Lower Bathonian of Cross Hands Quarry, Warwickshire represents a large, early basal tetanuran that is distinct from Megalosaurus and SDM 44.19 (see below). The specimens (WARMS G15770, G15771) preserve at least one autapomorphy as well as numerous features differentiating Cruxicheiros from coeval forms, but it is sufficiently incomplete that its placement within Tetanurae is uncertain (Benson & Radley 2010). Embasaurus minax Riabinin, 1931. Based only on a pair of dorsal vertebral centra from the Neocomian (Lower Cretaceous) of Kazakhstan, Embasaurus minax was originally assigned to Megalosauridae (Riabinin 1931). Like Becklespinax, the absence of pneumatic foramina (pleurocoels) in the posterior dorsal centra seems to exclude this taxon from Carcharodontosauria. The centra are amphiplatyan but the neural arches are not preserved. Embasaurus is here considered a nomen dubium, and there is little additional information to identify it more specifically than as Theropoda indet.


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Erectopus superbus (Sauvage, 1882) Huene, 1923. The partial skeleton forming the holotype of Megalosaurus superbus was collected from the lower Albian (Douvilleiceras mammilatum Zone) phosphatic ‘La Penthiève Beds’ of Bois de la Penthière, near Louppy-le-Château, Meuse, France (Sauvage 1882; Huene 1923; Allain 2005b). The material pertains to a medium-sized theropod and consists of a partial maxilla, incomplete manus, left femur and tibia, left calcaneum and right metatarsal II (Allain 2005b). Huene (1923) placed the species into a new genus, Erectopus, based on the morphology of the hind limb elements. The anteromedial (palatal) process of the maxilla is located ventrally, adjacent to the dorsal margins of the paradental plates, as in Marshosaurus and Sinraptor. The femur suggests avetheropod affinities as it lacks a proximal articular groove (Hutchinson 2001; Benson 2010a), and the condyles are separated by a shallow anteroposteriorly oriented trough on the distal surface. The distinct, step-like supraastragalar buttress of the tibia is distinct from the reduced buttress of carcharodontosaurians. We suggest that Erectopus represents a noncarcharodontosaurian allosauroid, possibly a metriacanthosaurid. Gasosaurus constructus Dong, 1985. Gasosaurus is based on vertebrae, a humerus, and highly abraded and reconstructed pelvic and hind limb material from the Middle Jurassic Xiashaximiao Formation of Dashanpu, Sichuan, China and was originally described as a megalosaurid. Holtz (2000) recovered Gasosaurus as a basal coelurosaur but Currie (in Holtz et al. 2004) stated that new specimens indicated it was a primitive ‘carnosaur’ (allosauroid). Many potentially informative features of the holotype skeleton (IVPP V7265) are difficult to assess based on published descriptions and images. The taxon represents a tetanuran based on the presence of a pubic peduncle of the ilium that is substantially larger than the ischial peduncle. Examination of casts reveals that the lesser trochanter does not rise above the level of the femoral head (contra Holtz 2000; Holtz et al. 2004). Instead, the proximal portion of the femoral head is broken and the lesser trochanter reaches approximately midlevel of the head as in non-coelurosaurian tetanurans. For now Gasosaurus is best regarded as having an uncertain position within Tetanurae and probably outside Coelurosauria; however, detailed restudy of the holotype is underway (D. Hone pers. comm.). Iliosuchus incognitus Huene, 1932. Iliosuchus includes a small partial ilium from Stonesfield, the topotype locality of Megalosaurus bucklandii (NHMUK R83; Taynton Limestone Formation, Bathonian, Middle Jurassic, England) and two referred ilia (OUMNH J.29780 [Galton 1976] and OUMNH J.29871 [Foster & Chure 2000]). The presence of a swollen ridge on the lateral surface of the ilium has been considered the diagnostic feature of this taxon. It has been used to unite the genus with the basal tyrannosauroid

Stokesosaurus clevelandi from the Late Jurassic of North America (Galton 1976), but this feature has a wider distribution among theropods. Benson (2009b) observed that it was also present in M. bucklandii, suggesting that I. incognitus showed no diagnostic features and should be considered a nomen dubium, representing either an indeterminate avetheropod or a juvenile specimen of M. bucklandii. OUMNH J.29871 is distinct from the holotype of I. incognitus because it has a vertical, not posterodorsally inclined, median ridge, larger size, and several small ‘accessory’ ridges anteriorly and posteriorly on the lateral surface. However, it cannot be identified other than as Tetanurae indet. Kaijiangosaurus lini He, 1984. Seven cervical vertebrae constitute the holotype of Kaijiangosaurus lini (CCG 20020), which was discovered in the Middle Jurassic Xiashaximiao Formation of Sichuan Province, China (He 1984). These elements are not elongate and have a flat anterior surface and a concave posterior surface, which are not significantly offset from one another. The neural spine is anteroposteriorly short and slightly posteriorly inclined. These features suggest placement as a basal tetanuran or basal averostran. Referred specimens include a tooth, jugal, two dorsal and seven caudal vertebrae, scapula and coracoid, humerus, proximal ulna, partial manus, femur, partial tibia and fibula, tarsus, and incomplete pes. Unfortunately it is not clear whether they collectively represent a single taxon. Restudy of these materials is needed. Kelmayisaurus petrolicus Dong, 1973. A left dentary and maxilla constitute the holotype (IVPP V 4022) of this taxon, which was recently redescribed, diagnosed, and assigned to Carcharodontosauridae (Brusatte et al. 2011). The presence of Kelmayisaurus in the Lianmugin Formation (Early Cretaceous) of China emphasizes the lengthy tenure of carcharodontosaurids in Asia. Orkoraptor burkei Novas, 2008. Orkoraptor is based on fragmentary remains from the Pari Aike Formation (?Maastrichtian) near Los Hornos Hill, Santa Cruz Province, Argentina. These were originally identified as a coelurosaur but have now been demonstrated to pertain to a megaraptoran allosauroid (Benson et al. 2010) based on the presence of proximal caudal pleurocoels, neural arch foramina and laminae and an Aerosteon-like postorbital. The affinities of Orkoraptor within Megaraptora remain unresolved due to the incomplete nature of the holotype and only specimen (MPM-Pv 3457). Poekilopleuron schmidti Kiprianow, 1883. This taxon is based on a three partial ribs and a distal humerus from the Albian–Cenomanian Sekmenevsk Formation of Tuskar, near Meshkovo, in western Russia. Interestingly, Kiprianow (1883) presented several histological sections of these bones showing details of the internal microstructure, one



of the earliest examples for dinosaurs. The bones themselves are quite fragmentary and largely uninformative. The humerus seems to exhibit a hollow interior (also present in small ornithopods, but these are smaller than P. schmidti) and the distal end is flattened anteroposteriorly. The radial condyle and ectepicondyle are broken but the ulnar condyle is intact and wraps slightly onto the posterior surface. There is a small entepicondyle on the medial side. At present we can only tentatively confirm the theropod nature of this element, and the species cannot be assigned to the genus Poekilopleuron. Rapator ornitholestoides Huene, 1932. The holotype of Rapator comprises only a left metacarpal I (NHMUK R3718), from the Griman Creek Formation (Albian) of Lightning Ridge, Australia. It was originally considered to pertain to a coelurosaur similar to Ornitholestes (Huene 1932), and more recently as manual phalanx I-1 from an alvarezsaurid (Holtz et al. 2004). However, Agnolin et al. (2010) noted similarities with metacarpal I of Australovenator and Megaraptor, and suggested that Rapator was a nomen dubium belonging to Megaraptora. We agree with this morphological assessment, but in the absence of comparison to the same element in Neovenator (currently unknown), we cannot place it more specifically than Neovenatoridae indet. Siamosaurus sutheethorni Buffetaut & Ingavat, 1986. Siamosaurus is based on isolated teeth (TF 2043a-i) from the Sao Khua Formation (Barremian–Aptian) of Phu Wiang, Thailand that resemble those of spinosaurids in exhibiting fluted enamel, a less recurved profile and relatively rounded cross section (although much less so than in some spinosaurids). The teeth do resemble those of spinosaurids but not enough material is known to support retention of a specific taxon, and their theropod affinities have been questioned (e.g. Sues et al. 2002; Holtz et al. 2004). They are here referred to as ?Spinosauridae indet., but we note that a partial skeleton from the Aptian Khok Kruat Formation of Thailand may eventually permit a more secure identification (Buffetaut et al. 2004, 2005; Milner et al. 2007). ‘Streptospondylus’ cuvieri Owen, 1842. Owen (1842) based Streptospondylus cuvieri on a partial dorsal vertebra, neural spine and tooth from the Inferior Oolite Formation (Aalenian–Bajocian) in the collection of Mr Kingdon of Chipping Norton, Oxfordshire, England. These were associated with a “broad flat bone” and fragments of long bones (Owen 1842, p. 90). It was subsequently transferred to Megalosaurus by Huene (1908, 1932) as the new combination M. cuvieri. Notably, the holotype specimens of Streptospondylus altdorfensis, Eustreptospondylus and Piveteausaurus were all at one time referred to S. cuvieri (Nopcsa 1906; Piveteau 1923; Huene 1932). Unfortunately,

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the holotype and referred specimens of S. cuvieri were never figured and are now presumed lost. The holotype vertebra is likely to have been an anterior dorsal because “the articular surfaces of the ribs are, as usual, close to the anterior part of the body of the vertebra”, which bore a convex anterior face (Owen 1842, p. 88). The centrum also had a large, single pneumatic opening that deeply invaginated into the body of the bone. Typical laminae were present on the base of the neural arch. The neural spine exhibited pronounced attachments for the interspinous ligaments along its anterior and posterior edges. The tooth was described as conical and hollow, and may not pertain to a theropod. The remaining specimens indicate an indeterminate tetanuran theropod that cannot be referred to the genus Streptospondylus. Owen also mentioned a second specimen, a posterior dorsal vertebra from the “jet-rock (lias shales)” (Jet Rock Formation, Lower Toarcian; Howarth 1980) of Whitby in the collection of Mr Ripley, as “referable to the present genus” (Owen 1842, p. 90). It too is lost but was apparently similar in basic morphology to the type specimens of ‘S.’ cuvieri, although more complete. It was clearly opisthocoelous and preserved more of the neural arch including the zygapophyses (“oblique processes”; Owen 1842, p. 90). The presence of a large, single pneumatic foramen in the centrum is a synapomorphy of Tetanurae and the Toarcian specimens may be one of the earliest records of the clade, but as no diagnostic features were described they remain Tetanurae indet.

Suchosaurus cultridens (Owen, 1841) Owen, 1842. This taxon was originally named Crocodilus cultridens for a single tooth (NHM R36536) that Owen (1841 in Owen 1840–1845) thought belonged to a crocodilian; he later removed it to the new genus Suchosaurus (Owen 1842). Only much later, following the discovery of Baryonyx, was the material re-examined and referred to the Baryonychinae (Buffetaut 2007, 2010; Mateus et al. 2011). The tooth itself is not diagnostic beyond the family level, and we consider it to be a nomen dubium.

Suchosaurus girardi Sauvage, 1897–1898. The material on which Sauvage based Suchosaurus girardi is slightly more complete than that used by Owen to describe the type species, S. cultridens, and includes a jaw fragment with teeth. These specimens bear several features in common with baryonychines (Buffetaut 2007, 2010; Mateus et al. 2011) but no diagnostic qualities of their own. S. girardi is therefore a nomen dubium; it cannot be formally referred to the later-named genus Baryonyx without subsuming that name into Suchosaurus. However, it derives from the Lower Barremian Papo Seco Formation of Portugal, where Baryonyx is known from other material (Mateus et al. 2011) and may represent the same taxon.

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Szechuanosaurus campi Young, 1942. Young (1942) named this theropod taxon based on four isolated teeth (IVPP V235, 236, 238, 239) from the Late Jurassic Kuangyuan Formation of Sichuan Province, China, but these are not diagnostic below the level of Theropoda. As this is the type species of the genus Szechuanosaurus, that taxon cannot be diagnosed and should not be used to receive additional species. Unquillosaurus ceibalii Powell, 1979. Based on an isolated pubis from the Los Blanquitos Formation (late Santonian–?Campanian) of Arroyo El Morterito (Campanian), Salta, Argentina, Unquillosaurus has been difficult to place phylogenetically. Originally described as the left pubis of an indeterminate large theropod (Powell 1979), it was recently redescribed as belonging to a giant maniraptoran (Novas & Agnolin 2004). In contrast to all previous workers, we are certain that this specimen (PVL 3670-11) is a right pubis. The ‘lateral crest’ (“cresta lateral”; Powell 1979, fig. 7) is the pubic apron, which bears a rugose contact surface for the opposing element at approximately midshaft. The obturator foramen, which is genuinely open, is flush with the lateral surface but inset from the medial surface at the proximal end. A fossa is present close to the contact for the ilium, which may represent the attachment area for M. ambiens (Carrano & Hutchinson 2002). In anterior view, the lateral margin is nearly straight but the medial margin is sinuous, suggesting the presence of a foramen between the pubes located proximal to the contacting distal ends. The distal end itself is eroded and broken, and lacks an anterior expansion although this may have been present when complete. It may also have been expanded considerably posteriorly; the posterior margin of the shaft forms an acute angle before terminating at a break. In all, the morphology of this bone resembles that of Giganotosaurus in many respects, and we suggest that it belongs to a carcharodontosaurid. It is possibly distinct enough to consider valid. Valdoraptor oweni (Lydekker, 1889) Olshevsky, 1991. Known only from left metatarsals II–IV (NHMUK R2559), this material, from the Tunbridge Wells Sand Formation (Valanginian) of Cuckfield, Sussex, was originally identified as a specimen of Hylaeosaurus (Owen 1858) and later named as a species of Megalosaurus (Lydekker 1889). Its age (Valanginian) alone makes it an unlikely member of this genus, and Olshevsky (1991) removed it to a new form, Valdoraptor. Although there may be other distinctive features about the holotype (Naish & Martill 2007), it is probably an avetheropod based on the trapezoidal cross section of metatarsal III, a unique, unambiguous synapomorphy in our analysis. Wakinosaurus satoi Okazaki, 1992. This taxon is based on a single tooth (KMNH VP 000,0016) from the Sengoku

Formation (Hauterivian–Barremian) of Fukuoka, Japan, that was distinguished by the presence of longitudinal striations. Unlike other teeth with similar features, it is strongly laterally compressed. Okazaki (1992, p. 88) referred Wakinosaurus to Megalosauridae and suggested that it might share affinities with the tooth taxon Prodeinodon kwangshiensis. Although the tooth is distinctive, it cannot be assigned either to Megalosauridae or to any particular clade within Theropoda at this time.

Species previously assigned to the genus Megalosaurus A host of species ranging in age from Late Triassic through Late Cretaceous has been placed within the genus Megalosaurus, but presently we consider only the type species M. bucklandii to be properly assigned to it (Benson 2009a, 2010a). All other taxa originally referred to Megalosaurus that are represented by diagnostic materials have since been assigned to other genera. They have been included in our analysis or discussed in the previous section and are only listed here: Carcharodontosaurus saharicus, Dilophosaurus wetherilli, Duriavenator hesperis, Erectopus superbus, Magnosaurus nethercombensis, Majungasaurus crenatissimus, Proceratosaurus bradleyi and Valdoraptor oweni. Betasuchus bredai is considered to belong to Ceratosauria (Carrano & Sampson 2008). A number of theropod taxa have been included within Megalosaurus in previous taxonomic revisions although they were not originally named as species of that genus. These include Antrodemus valens (Nopcsa 1901), Ceratosaurus nasicornis (Cope 1892), Deinodon horridus (Nopcsa 1901), Dryptosaurus aquilunguis (as a subgenus of Megalosaurus; Depéret & Savornin 1928), Iliosuchus incognitus (Kuhn 1939; Romer 1956), Nuthetes destructor (Romer 1956), Poekilopleuron bucklandii (as Megalosaurus poikilopleuron; Huene 1926a) and Torvosaurus tanneri (Paul 1988a). Laelaps trihedrodon Cope, 1877 has been previously recombined as Dryptosaurus trihedrodon (Cope 1878), Creosaurus trigonodon (Osborn 1931) and Megalosaurus trihedrodon (Nopcsa 1901). It is based on a partial dentary (AMNH, lost specimen) and referred teeth (AMNH 5780) from the Morrison Formation (Kimmeridgian–Tithonian) of Cañon City, Colorado that can be referred to Allosaurus fragilis (Chure 2001b; Holtz et al. 2004). Several valid or potentially valid forms are based on specimens that have at times been assigned to Megalosaurus but not as distinct species: Becklespinax altispinax (as M. dunkeri), Eustreptospondylus oxoniensis and Streptospondylus altdorfensis (both as M. cuvieri). In addition, Dakosaurus maximus Quenstedt, 1858 is a valid metriorhynchoid taxon originally based on a dentary fragment from the Kimmeridgian–Tithonian of Schnaitheim, Germany (Megalosaurus sp. in Quenstedt, 1843).



Other species were erected without a description, diagnosis or indication of type material and therefore represent nomina nuda (Article 12; ICZN): M. cachuensis from the Middle Jurassic Dapuka Group of Xinjiang Uygur Zizhiqu, China (Zhao in Weishampel et al. 2004b), possibly a misspelling of M. dapukaensis Zhao, 1986 (also a nomen nudum; Olshevsky 1991); and Megalosaurus tibetensis Zhao, 1986. The remaining species, mostly fragmentary and non-diagnostic, are assigned as follows and listed alphabetically by species. Zanclodon cambrensis Newton, 1899. This species is based on a natural mould of a left dentary (NHMUK R2912) from the Rhaetian beds near Bridgend, Wales. Kuhn (1939, 1965) listed it as Zanclodon (Megalosaurus) cambrensis, probably indicating that he believed an earlier referral to Megalosaurus existed. Waldman (1974) and Molnar (1990) considered that the holotype bore detailed resemblance to ‘Megalosaurus’ (now Duriavenator) hesperis and M. bucklandii; Galton (1998, 2005) transferred the taxon to Megalosaurus cambrensis based on Waldman’s (1974) observations. However, NHMUK R2912 is substantially different from Megalosaurus, indeed from tetanurans generally, and is an indeterminate theropod outside Averostra or a more basal predatory archosaur (Rauhut & Hungerbühler 2000; Naish & Martill 2007; Benson 2010b). Megalosaurus chubutensis del Corro, 1974. M. chubutensis is based on a single lateral tooth (MACN 18.189) from the upper member of the Cenomanian (Cerro) Castillo Formation north of Cerro Crettón, Chubut, Argentina (del Corro 1974; Bridge et al. 2000). It is poorly preserved but moderately large (85 mm long), bearing very fine posterior serrations (approx. 2 per mm) that reach the tooth base. The anterior carina is strongly recurved but the posterior is nearly straight. In posterior view (del Corro 1974, fig. 1) the posterior carina exhibits a distinct apicomedial curvature. The tooth resembles those of abelisaurids but is extremely large compared to known forms. Megalosaurus cloacinus Quenstedt, 1858. This is among the geologically oldest species referred to the genus Megalosaurus as the holotype and only specimen (SMNS 52457) was found in the Rhät bonebed (Rhaetian–early Hettangian) of Baden-Württemburg, Germany. Huene (1932) referred the holotype dentary of Zanclodon cambrensis to M. cloacinus (as Gresslyosaurus (?) cloacinus) based on unspecified dental similarity. However, SMNS 52457 is a serrated, recurved tooth of the form typical for theropods. It is mesiodistally slender but does not show any diagnostic features and is therefore Theropoda indet.; additional referrals to M. cloacinus cannot be made with confidence.

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Megalosaurus dunkeri Dames, 1884. This taxon is founded on a worn tooth from the ‘lower Wealden’ (Early Cretaceous) of Nordrhein-Westphalen, Germany (University of Marburg N 84; Dames 1884; Huene 1923). Lydekker (1888, pp. 165–168) subsequently referred a large number of NHMUK specimens from the Lower Cretaceous of England to M. dunkeri although he noted that not all of these specimens lacked anterior serrations, a supposedly diagnostic feature. Lydekker (1890a) later referred ‘most or all’ of the specimens from Cuckfield, Sussex and the ‘Weald Clay’ of the Isle of Wight to Megalosaurus oweni for unspecified reasons. Huene (1923) also considered that the holotype vertebrae of Becklespinax altispinax might be referred to M. dunkeri (q.v.). The holotype of M. dunkeri does not appear diagnostic beyond the level of Theropoda and referred specimens cannot reliably be assigned to the species. Megalosaurus hungaricus Nopcsa, 1902. Nopcsa (1902, figs 1–6) described and illustrated a single tooth (MAFI Ob 3106) from the Late Cretaceous coal-bearing strata of Nagy-Bároth, Romania (then part of the AustrianHungarian Empire) as the holotype of M. hungaricus. Nopcsa considered this stratum equivalent to the Maastrichtian beds at Szentpéterfalva but the proximity of this locality to the Early Cretaceous Brusturi site suggests caution in the absence of detailed geological reconnaissance. The tooth is moderate in size (c.22 mm long). The denticles on the posterior carina show a distinctive, possibly autapomorphic morphology: they are separated by spaces equivalent to the proximodistal width of a single denticle (Nopcsa 1902, p. 104). The anterior denticles are slightly smaller than the posterior ones, with a denticle size difference index around 1.25 (Nopcsa 1902), similar to those of some tyrannosauroids and dromaeosaurids (Rauhut & Werner 1995; Sweetman 2004). M. hungaricus teeth are less transversely compressed than those from the same general age in Europe referred to M. pannoniensis (Nopcsa 1902). Nonetheless, the latter has been considered a senior synonym of M. hungaricus (Lapparent 1947; Lapparent & Zbyszewski 1957). Huene (1926a, b) noted that ‘M.’ hungaricus could not be assigned to a genus and we concur that the taxon should be considered Theropoda indet., although it is possible that they represent either tyrannosauroids or dromaeosaurids. Megalosaurus inexpectatus del Corro, 1966. This second species of Megalosaurus from the Chubut Group was based on five large teeth (holotype, MACN 18.172) associated with the holotype skeleton of the sauropod Chubutisaurus insignis (del Corro 1975). They derived from the Albian–Cenomanian Bayo Overo Member of the Cerro Barcino Formation at Paso de Indios, Argentina (del Corro 1966). At least two of the teeth (del Corro 1966, pl. 1, figs 1, 2) exhibit distinct banding of the enamel as is common

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among tetanurans (Brusatte et al. 2007). The holotype shows an overall curvature and apical wear reminiscent of carcharodontosaurid teeth, to which it should be compared. Megalosaurus ingens Janensch, 1920. This taxon consists of a large tooth (MB.R1050; 15 cm long) from Tendaguru Quarry B in the Upper Dinosaur Member of Tanzania (Tithonian; Janensch 1920, 1925). Both anterior and posterior carinae are coarsely serrated (≤1 per mm) along their entire lengths. The enamel shows transverse bands that are particularly pronounced posteriorly as in many tetanurans (Brusatte et al. 2007). Numerous other teeth (25 MB specimens; NHMUK R5556, R5557, R6758), some from other levels of the Tendaguru Beds, have been referred to this taxon based primarily on the coarseness of the serrations and their large overall size (Janensch 1925). We do not consider these characteristics sufficient to define a taxon or refer additional specimens, and refer M. ingens to Tetanurae indet. Megalosaurus insignis Eudes-Deslongchamps & Lennier in Lennier, 1870. The holotype tooth of M. insignis was found in the lower Kimmeridgian deposits of Cap de la Hève, France (Valenciennes 1863; Lennier 1870; Huene 1932). It was described as large (80 mm long), laterally compressed, recurved, and serrated along the posterior carina but was missing its apex (Lennier 1870, 1887, pl. 1, figs 1–3). It is indeterminate below the level of Theropoda. Additional specimens include a pedal phalanx and ungual from the same area (Lennier 1887; Huene 1932) but both appear to pertain to sauropods (Lennier 1887, pl. 1, figs 4–7). Teeth, vertebrae, claws, an ulna and partial femur from the Kimmeridgian of Portugal were later assigned to M. insignis (Lapparent & Zbyszewski 1957); although some elements preserve interesting morphological details, placement within this taxon is not justified. The Portuguese collection includes an articulated series of five anterior caudal vertebrae from Praia da Areia Branca that are associated with two additional articulated caudals. The transverse processes are upturned and extend far laterally whereas the centra lack neural arch laminae or centrum pleurocoelous fossae and bear tall, vertical neural spines. They may belong to a teleosaurian crocodilian (Chabli 1986). Megalosaurus lonzeensis Dollo, 1883. This species was founded on a partial manual ungual from the Santonian ‘glauconite argileux’ of Lonzée, Namur, Belgium (Dollo 1883). The small size, mediolaterally narrow dimensions and details of the vascular traces more closely resemble the manual unguals of coelurosaurs. Huene (1926a, 1932) placed it among ornithomimids but it is best referred to as Coelurosauria indet.

Megalosaurus lydekkeri Huene, 1926a. A tooth from the Lower Lias Group of Lyme Regis (NHMUK 41352) may correspond to reports of Megalosaurus (Dawkins in Huxley 1870) or M. bucklandii (Phillips 1871) from the Lias of Dorset. It was first formally recorded as Zanclodon sp. b (Lydekker 1888) and later questionably assigned to Megalosaurus sp. (Huene 1908), ‘Zanclodon’(?) sp. (Huene 1926a, p. 36) and ‘Megalosaurus’ (gen.?) lydekkeri n. sp. (Huene 1926a, table 1). The crown is laterally compressed and smoothly curved throughout its length, bearing nearly fully serrated anterior and posterior carina and faint longitudinal striations. The latter may suggest non-tetanuran affinities, because the tooth lacks the specialized features of tetanurans in which such striations also occur (e.g. spinosaurids). Otherwise the specimen is indeterminate within Theropoda. Megalosaurus meriani Greppin, 1870. This fragmentary form derives from the ‘Virgulian’ Basse Montagne quarry near Moutier, Kanton Bern, Switzerland, now known to belong to the lower Reuchenette Formation (early Kimmeridgian). It was originally described as ‘a complete skeleton’, ‘a large part’ of which was in the Musée de Bâle (now the Naturhistorisches Museum Basel). However, the accompanying description and plate (Greppin 1870, pl. 1, figs 2–5) demonstrate that the majority of the specimens pertain to a sauropod; these were later removed from M. meriani and now constitute the holotype of Cetiosauriscus greppini (Huene 1927; Meyer & Thuring 2003). The holotype of M. meriani is therefore restricted to the only demonstrably theropod element, a single tooth (MH 350; Greppin 1870, pl. 1, fig. 1). It is mediolaterally thick, bears longitudinal striations on the internal face and fine serrations on both carinae. Overall it resembles the anterior teeth of Ceratosaurus (Madsen & Welles 2000), although we assign it to Ceratosauria indet. rather than this genus. Megalosaurus mersensis Lapparent, 1955. M. mersensis was based on a partially articulated series of 23 vertebrae from the Bathonian El Mers Formation north of Tizi n’Juillerh, Morocco (Lapparent 1955). These are not well preserved but represent much of the neck and back as well as parts of the sacrum and tail. The axis lacks pneumatic foramina and bears only a small diapophysis. It is difficult to determine from the illustrated vertebrae (Lapparent 1955, pl. 3) whether any of the subsequent vertebral centra are pneumatized. The cervical centra appear to have only weakly offset faces and are amphicoelous (Chabli 1986). The anterior dorsal (Lapparent 1955, pl. 3, fig. 6) has a horizontally oriented transverse process and a neural spine at least as tall as the height of the centrum. The centra of two succeeding dorsals (Lapparent 1955, pl. 3, fig. 8) are slightly longer than tall and have a shallow ventral arch. The vertebrae are quite different from those of most megalosauroids (e.g. Baryonyx, Eustreptospondylus, Piatnitzkysaurus) in

The phylogeny of Tetanurae one or more of these features. Without additional study of the type specimen we cannot determine the validity of the species ‘M.’ mersensis, although we agree that it does not belong within the genus Megalosaurus and instead seems likely to represent a teleosaurid mesosuchian (Chabli 1986).


Saurocephalus monasterii Munster, 1836. Münster ¨ (1836, pl. 3, fig. 15a–d) erected a new species of the fish genus Saurocephalus, S. monasterii based on a recurved, serrated tooth from the Oxfordian Korallenkalk of the Lindner Berge, Hanover, Germany. Windolf (1997) transferred the taxon to Megalosaurus but Münster’s tooth cannot be identified past the level of Theropoda indet. Aggiosaurus nicaeensis Ambayrac, 1913b. A partial mandible from the upper Oxfordian in the region of Cap d’Aggio-La Turbie, Monaco was reported as a plesiosaur or teleosaurid crocodile by Depéret (in Ambayrac 1913a) and made the holotype of the crocodilian taxon Aggiosaurus nicaeensis (Ambayrac 1913b). Subsequently the material was considered to represent a carnosaurian (Huene 1932) and transferred to Megalosaurus nicaeensis (Romer 1956; Steel 1970). The teeth of A. nicaeensis are conical and only slightly recurved, with fine enamel ridges; replacement teeth are present inside the hollow roots of functional teeth. Therefore, A. nicaeensis can be placed within Metriorhynchidae, although as a nomen dubium (Buffetaut 1982). Megalosaurus obtusus Henry, 1876. The holotype specimens of this form were recovered from Rhaetian marine outcrops near Moissey, Franche-Comte, France. The four referred teeth are fragmented and poorly preserved but show distinct carinae and serrations (Henry 1876, pl. 1, figs 1–4). Thus they should not be referred to Plateosaurus (Huene 1908) but as indeterminate specimens within Theropoda or another predatory archosaur clade. Megalosaurus pannoniensis Seeley, 1881. The Maastrichtian Gosau Formation near Weiner Neustadt, Austria produced two teeth (GMUV uncatalogued) that Seeley (1881) referred to M. pannoniensis (for which M. pannonicus is a lapsus calami; Huene 1926a). Huene (1926a) considered them unlikely to pertain to Megalosaurus due to their young geological age and noted that the illustrated specimen “does not show anything concerning the nature of the genus” (Huene 1926a, p. 81). The serrations are fine, ∼ 3.5 per mm on the anterior carina and ∼ 2 per mm on the posterior carina. The anterior serrations terminate before the base of the tooth and Seeley (1881, p. 670) noted the presence of transverse “lines of growth” across the enamel that are now regarded as enamel ‘wrinkles.’ Together these features allow assignment to Tetanurae indet., and the relatively small anterior denticles suggest possible assignment

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to either Dromaesauridae or Tyrannosauroidea (Rauhut & Werner 1995; Sweetman 2004). Megalosaurus pombali Lapparent & Zbyszewski, 1957. This species was named for three teeth from unnamed Callovian–Oxfordian beds near Pombal, Portugal. Lapparent & Zbyszewski (1957) distinguished it from M. insignis by the larger size and thicker cross sectional dimensions. Only the apical one-third of the anterior carina is serrated. The figured tooth (Lapparent & Zbyszewski 1957, pl. 28, fig. 105) is indistinct from other theropod teeth and should be relegated to Theropoda indet. However, a set of eight vertebrae from Porto das Barcas was also referred to M. pombali. Although poorly described and figured and not referable to M. pombali, they warrant further study to determine whether they represent a diagnosable taxon. Massospondylus rawesi Lydekker, 1890b. The holotype tooth, from the Takli Formation (late Maastrichtian; Sahni et al. 1984) of Maharashtra, India, is recurved, serrated and laterally compressed (NHMUK R4190). Although the species has been referred to Megalosaurus (Vianey Liaud et al. 1987) and Orthogoniosaurus (Olshevsky 1991), it is not diagnostic past the level of Theropoda indet. The fine serrations and stout proportions resemble the condition in abelisaurids (Carrano et al. 2010). Megalosaurus schnaitheimii Bunzel, 1871. This taxon is based on teeth originally described and illustrated as Megalosaurus sp. by Quenstedt (1852). They derive from Late Jurassic marine deposits (possibly the Korallenkalk) of Schnaitheim, near Brenz, Germany, and may pertain to Dakosaurus maximus. Zanclodon silesiacus Jaekel, 1910. A tooth from the Muschelkalk of Gogolin, Oberschlesien, in the collection of the University of Griefswalden/Göttinger is the holotype of Zanclodon silesiacus Jaekel, 1910. This specimen was referred to ‘Megalosaurus’ silesiacus by Kuhn (1965). It has the recurved, serrated form typical of theropods and could be considered as Theropoda indet., but we cannot rule out the possibility that it represents a ‘rauisuchian’ archosaur. Megalosaurus terquemi Gervais, 1859. This species is based on three isolated teeth from the Hettangian ‘Angulatus Beds’ of Hettingen, near Moselle, Germany (Gervais 1853; Terquem 1855; Huene 1926a). Huene considered them sufficiently distinct to refer to them as ‘Megalosaurus’ (gen. 2) terquemi (Huene 1926a, p. 80) or Megalosauridorum gen. indet. terquemi (Huene 1932 p. 219), noting that the teeth were longer, sharper and more lingually compressed than those of Plateosaurus (Huene 1932). Buffetaut et al. (1991b) considered that the teeth belonged to phytosaurs. As they were housed


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at the Muséum du Caen, we presume they were lost when that institution was destroyed by incendiary bombs during the Second World War, although they were not specifically listed as such (Bigot 1945). We refer them to Archosauria indet. Finally, the tendency to assign tracks to Megalosauridae or Megalosaurus is largely without a firm anatomical basis, and all such occurrences should be considered Theropoda indet. This is particularly true of large theropod trackways from the Early Cretaceous of England (Delair 1982) and Spain (Sanz et al. 1990), which are far removed temporally from any European occurrences of this family. It is beyond the scope of this paper to review all the individual specimens assigned to the genus Megalosaurus, but we are doubtful that any (aside from those specifically reviewed above and in Benson 2010a) have been correctly attributed.

Fragmentary occurrences Africa. The Late Jurassic–?earliest Cretaceous Tendaguru Formation includes a small number of theropod specimens compared to the penecontemporaneous Morrison deposits of North America, but probably includes 4–5 taxa nonetheless. Among them are a slender ceratosaur, Elaphrosaurus; a larger, more robust relative that includes some of the material referred to Ceratosaurus? roechlingi and several hind limb bones (Janensch 1925; Carrano et al. 2002; Carrano & Sampson 2008); and at least one small abelisauroid (Rauhut 2005b). Additionally, Janensch (1925) named Allosaurus? tendagurensis based on a large, damaged tibia (see above). Rauhut (2005b) also identified two other tibiae, one small, as indeterminate basal tetanuran remains. An isolated right ilium (MB.R 3628 = St 233) also derives from a tetanuran (Fig. 11) but is unusual in exhibiting a very widely open preacetabular notch, a strongly but broadly downcurved dorsal margin of the preacetabulum, and a markedly tapering postacetabular process. In addition, it has a preac-

Figure 11. Possible Late Jurassic carcharodontosaurian ilium (MB.R 3628 = St 233) from Tendaguru, Tanzania. Abbreviations: ac, acetabulum; ip, ischial peduncle; pn, preacetabular notch; pp, pubic peduncle. Scale bar = 10 cm.

etabular fossa (an avetheropodan feature), a pubic peduncle with length:width proportions similar to allosaurians (∼ 2.0) but shorter than coelurosaurs (∼ 3.0) and longer than all non-avetheropods and metriacanthosaurids (≤1.7), and a ventrally rather than anteroventrally oriented pubic peduncle as in carcharodontosaurians. Taken together, these features suggest that the ilium may pertain to a carcharodontosaurian. The many isolated Tendaguru teeth are largely Theropoda indet., although some may pertain to ceratosaurians as they possess longitudinal striations on the lingual surface similar to those on the premaxillary teeth of Ceratosaurus (Madsen & Welles 2000). Approximately contemporaneous with the Tendaguru materials, theropod teeth from the Late Jurassic of Ethiopia have been described as cf. Acrocanthosaurus sp. (Goodwin et al. 1999). However, this identification was based solely on the presence of ‘chisel-shaped’ denticles, a feature present in, but not unique to, Acrocanthosaurus. One specimen displays surface crenulations (‘wrinkled enamel’), but these are widely distributed among tetanurans (Brusatte et al. 2007) and we assign these teeth to Tetanurae indet. The more southerly exposures of the ?Late Jurassic Kadzi Formation of Zimbabwe have also produced theropod remains, including femora referred to Allosauridae? indet. (Raath & McIntosh 1987). These femora are badly damaged but exhibit several features that suggest affinities with Ceratosauria, rather than Tetanurae. They are currently under study (E. Roberts, P. O’Connor & M. Carrano in prep.). A vertebral centrum from the Hauterivian–Barremian Cabao Formation of Libya was referred to ?Spinosaurus sp. (El-Zouki 1980; Tawadros 2001). The proportions match well with the posterior dorsals of Baryonyx and Suchomimus but are relatively shorter than those of Megalosaurus and carcharodontosaurids. The (presumed) ventral surface is only slightly narrower than the centrum, similar to the condition in Megalosaurus and Suchomimus (El-Zouki 1980, pl. 2). We consider it likely that this specimen belongs to the clade Megalosauria. The tooth listed as cf. Baryonyx sp. from the same formation (Le Loeuff et al. 2010) is assigned here to Baryonychinae indet. A proximal femur (AM 6041) from the Berriasian–Valanginian Kirkwood Formation (Forster et al. 2009) of South Africa indicates the presence of a small tetanuran in these deposits. Forster et al. (2009) proposed that the inferred presence of a weakly anteromedially (rather than strictly medially) oriented femoral head suggested non-avetheropodan affinities. However, the femoral head of some allosauroids is oriented slightly (Allosaurus, Sinraptor; Benson 2009b, 2010a) or strongly (Neovenator; Brusatte et al. 2008) anteromedially and both the absence of a proximal articular groove and the presence of a well-developed accessory trochanter


The phylogeny of Tetanurae indicate avetheropodan affinities. We regard this specimen as Allosauroidea indet. but note that the head lacks the proximal inclination of most carcharodontosaurians. Spinosaurid and carcharodontosaurid remains, mostly teeth but also vertebrae (Rauhut 1999), have been found across North Africa in deposits ranging in age from Albian through Cenomanian (e.g. Bouaziz et al. 1988; Russell 1996; Rauhut 1999; Sereno et al. 2004). It is not clear whether all these specimens can be accommodated by currently named taxa, but this issue is unlikely to be decided without more complete materials. Sereno & Brusatte (2008) referred postcranial specimens including a pelvis and sacrum (MNN GAD1-2) from the Elrhaz Formation (Aptian–Albian) of Niger to the abelisaurid Kryptops palaios. This material was collected in situ but the holotypic maxilla (MNN GAD1-1) was found eroded free of the rock some 15 metres distant. Several features of the pelvis and sacrum were described as ‘strikingly primitive’ for an abelisaurid, such as the presence of five sacral vertebrae, a reduced supraacetabular shelf that is not continuous with the lateral wall of the brevis fossa, an iliac pubic peduncle that is substantially larger than the ischial peduncle, and the relatively high and dorsoventrally short proportions of the ilium (Sereno & Brusatte 2008, p. 23). However, these features, which are present generally in tetanurans, are not present in even the most basal ceratosaurs and would therefore represent homoplasies in an abelisaurid. Both Ceratosaurus and Elaphrosaurus have six sacral vertebrae, a hypertrophied supraacetabular shelf that is continuous with the lateral wall of the brevis fossa, an iliac pubic peduncle that is only slightly larger than the ischial peduncle, and relatively long, low proportions of the iliac blade. Furthermore, the ischia of MNN GAD1-2 contact via a medial ‘apron’ as in many tetanurans but unlike the flat medial contact of more basal theropods, whereas the presence of a ventral fenestra between the pubic distal expansions similar to that of Ceratosaurus (Benson et al. 2009), as figured by Sereno & Brusatte (2008, fig. 7B), cannot be determined due to damage. Instead, a proximodistally elongate fenestra is present between the pubic shafts proximal to the distal expansion, as seen in tetanurans (Benson et al. 2009). The presence of a subrectangular fenestra between the fourth and fifth sacral neural spines (Sereno & Brusatte, 2008, fig. 7) is identical to that of Giganotosaurus (MUCPv-Ch 1), as is the presence of a peg-and-socket iliac-ischial articulation (a carcharodontosaurian synapomorphy). Taken together, we consider it unlikely that MNN GAD1-2 pertains to an abelisaurid or even a primitive ceratosaur, and suggest that is represents a carcharodontosaurid, most likely the sympatric Eocarcharia (currently only known from skull bones). Asia. Nessov (1995) described ‘coelurid’ and ‘megalosaurid’ teeth from the Callovian Balabansai Svita at Sarykamyshai in the Fergana Valley, Kyrgyzstan. These

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were recently redescribed (Averianov et al. 2005) as belonging to tetanurans with possible affinities to Dromaeosauridae. The presence of a short anterior carina was noted as a particular similarity with dromaeosaurids, but in fact this feature is quite common among basal tetanurans and is recovered here as the primitive condition for the clade. Other aspects of the size and shape of these teeth are also consistent with identification as a non-coelurosaurian tetanuran. Six teeth from the Callovian–Oxfordian Djaskoian Formation of Teete Creek, Yakutia, Russia were identified as Allosaurus sp. (PIN 4874/2; Kurzanov et al. 2003). They are large, recurved and serrated along both carina but cannot be identified more specifically than Theropoda. Likewise, Allosaurus sp. has been reported from unspecified Late Jurassic strata near Datong, in the Xinrong District of Shaanxi Province, China (Lü & Hu 1998). They include anteroposteriorly short dorsal vertebrae that appear to bear deep lateral fossae, ‘honeycomb’ internal structure described by Lü & Hu (1998, pl. 1) seems to be a marrow cavity rather than camellate pneumatic architecture. The overall morphology and cavernous lateral depressions are loosely comparable to Torvosaurus posterior dorsal vertebrae. However, the published images are difficult to interpret, so we consider them Tetanurae indet cf. Torvosaurus. Buffetaut & Suteethorn (2007) reported a metriacanthosaurid (‘sinraptorid’) tibia from the Late Jurassic–Early Cretaceous Phu Kradung Formation of Thailand. As reported by these authors, this specimen (SM 10) shows various detailed similarities with Chinese metriacanthosaurid taxa that support this identification. Early Cretaceous deposits at Tsanmakou in Shaanxi, China have yielded a fragmentary theropod skeleton (IVPP V969) that was assigned to Allosauridae (Young 1958). The material was only briefly described and illustrated but was compared favourably to Antrodemus valens (= Allosaurus fragilis); Young (1958, p. 235) suggested it might represent a new taxon. The scapula was described as being robust and having a somewhat expanded distal end. The fibula differs from those of spinosaurids and other megalosaurians in possessing a prominent, centrally placed medial fossa, and from those of allosauroids in exhibiting an obliquely oriented, rather than horizontal, proximal margin of this fossa. Unlike tetanurans generally, the iliofibularis tubercle is quite prominent. This material may represent a ceratosaurian but further study is required. We refer it to Averostra indet. Chure et al. (1999) described a tooth (MDM 341) from the late Cenomanian–early Turonian Jobu Formation (Mifune Group) of Kunamoto Prefecture, Japan, as a possible carcharodontosaurid, although they noted several differences with the teeth of currently known forms. Both the arcuate wrinkles and overall proportions are reminiscent of carcharodontosaurids but are now known to occur widely within Tetanurae (Brusatte et al. 2007).


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A middle caudal vertebra, fragmentary tibia and the distal end of a phalanx (IVPP V2756) from the Early Cretaceous Jehol Group of Shaihaitzu, Liaoning were assigned to Megalosauridae (Hu 1963). However, the tibia (the only illustrated element) bears a small, discretely rounded lateral condyle at the proximal end that strongly recalls the morphology of more derived theropods. It may pertain to a coelurosaurian. A fragmentary tibia from the Upper Cretaceous of Syria (Hooijer 1968) probably represents a tetanuran based on its described similarity to the same bones in Carcharodontosaurus and Erectopus. In particular, the proportions of the distal end are transversely expanded, comparable to the condition in tetanurans generally. Other theropod teeth from Jurassic and Cretaceous deposits of eastern Asia, often ascribed to megalosaurids (YNUGI 10003; KS 7001; Hasegawa et al. 1992; Nessov 1995; Lim et al. 2002) or allosaurids (KPE 8004, 8005; Park et al. 2000) cannot be diagnosed beyond Theropoda (e.g. Averianov et al. 2005). Those from South Asia (Ghevariya 1988; Sahni & Bajpai 1988) are similarly indeterminate (e.g. Bajpai & Prasad 2000), although many may ultimately pertain to abelisaurids. Australia and Antarctica. Aside from Cryolophosaurus and Australovenator, most theropod remains from these continents are quite fragmentary. The purported tetanuran caudal vertebra (WAM 96.5.1; Long & Cruickshank 1996) from the Birdrong Sandstone north of Kalbarri, Western Australia is too fragmentary to identify with even this level of specificity, and we consider it to be Theropoda indet. We likewise assign the ‘carnosaur’ pedal phalanx from the Cenomanian–Turonian Molecap Greensand (WAM 92.7.1; Long 1995) and the ungual from the Valanginian–Albian Wonthaggi Formation once referred to Megalosaurus (NMV P.10058; Woodward 1906; Huene 1926a; Rich & Rich 1989) to Theropoda indet. A theropod ulna (NMV P186076; Rich & Vickers-Rich 2003) from the Eumeralla Formation at Dinosaur Cove, Victoria (Aptian–Albian) was referred to cf. Megaraptor by Smith et al. (2008) and Megaraptora (Agnolin et al. 2010); we consider it to belong to an indeterminate neovenatorid (Benson et al. 2010). A distal tibia (MLP 89-XII-1-1) from the Coniacian–Santonian Hidden Lake Formation of James Ross Island was identified as Tetanurae indet. (Molnar et al. 1996). These authors noted similarities with basal tetanurans, particularly Piatnitzkysaurus, in the shapes of the medial malleolus, medial buttress and fibular flange. The distal end has a ratio of transverse width to anteroposterior thickness of 2.35, lower than that of Piatnitzkysaurus (2.80; MACN CH 895) and most other basal tetanurans (Benson 2010a, table 4), but it is possible that this value has been reduced by abrasion in MLP 89-XII-1-1. The bone is quite distinct from the ceratosaur condition and differs

from Cryolophosaurus in the contour of the medial edge. Similarities to Piatnitzkysaurus observed by Molnar et al. (1996) are shared with megalosauroids more generally, so we suggest that this bone pertains to a megalosauroid, the first identified from either continent, and one of the latest known. Perhaps the best known tetanuran specimen from Australia is an astragalus recovered from the Wonthaggi Formation at Eagle’s Nest, Victoria (NMV P150070) that was originally referred to Allosaurus sp. (Molnar et al. 1981). Since its description, its taxonomic identity has been much debated (Welles 1983; Molnar et al. 1985; Rich 1996; Chure 1998) and it was most recently assigned to Australovenator sp. (Hocknull et al. 2009) and Abelisauroidea (Agnolin et al. 2010). Despite the arguments of Agnolin. et al. (2010), we agree that this specimen pertains to a megaraptoran allosauroid, but given its temporal and geographical distance from the type material of Australovenator, and its resemblance to the astragalus of Fukuiraptor, we cannot exclude that it belongs to a distinct member of this clade (Benson et al. 2010). Europe. The theropod knee joint once considered part of the lectotype of Scelidosaurus harrisonii (NHMUK 39496; Newman 1968) shows several tetanuran characteristics (Carrano & Sampson 2004). The proximal morphology of the tibia resembles the condition in megalosauroids more so than ceratosaurs or coelophysoids. The fibula has a large but shallow, posteriorly open fossa on the medial surface as in some basal tetanurans and non-megalosaurian megalosauroids, but unlike the deep, centrally placed fossa of ceratosaurs or the narrow posterior incisure of coelophysoids. Finally, the distal femur has a vertically oriented tibiofibularis crest as in most tetanurans. This Liassic specimen may therefore be one of the earliest known tetanurans, but until more complete remains are found it is uncertain whether these features are synapomorphies permitting inclusion within Tetanurae, or symplesiomorphies that are transformed in Ceratosauria (Benson 2010b). The informal name ‘Saltriosaurus’ has been used to refer to a large theropod discovered in the Sinemurian Saltrio Formation, near Saltrio, Varese, Italy, in 1996 (Dal Sasso & Brillante 2001; Dal Sasso 2003). The specimen (MSNM V3664) is incomplete but includes a furcula, scapula, humerus, manual elements and parts of the hind limb. Enough is preserved to support the theropod affinities of ‘Saltriosaurus,’ but more specific assignment must await further preparation and description of the material. The widespread presence of a furcula amongst theropods, including coelophysoids (Tykoski et al. 2002; Rauhut 2003; Rinehart et al. 2007), demonstrates that this feature alone is insufficient to identify ‘Saltriosaurus’ as a tetanuran (Benson 2010b). Little can be said of the theropod vertebra described as Megalosaurus by Huene (1966) from a glacial erratic


The phylogeny of Tetanurae boulder in Ahrensburg, Schleswig-Holstein, Germany, which he considered to have been derived from the ‘Lias epsilon’ (Toarcian). Interestingly, the fragmentary specimen exhibits a convex anterior centrum face and a substantial lateral pleurocoelous fossa. It may therefore represent a mid-posterior dorsal from a large averostran theropod, although it could also represent a small sauropodomorph. The Jurassic theropods of Europe have a complex history. Reynolds (1939) assigned a collection of large theropod bones from the early Bathonian of New Park and Oakham Quarries to Megalosaurus. Benson (2009b) reviewed the New Park Quarry material and established that all bones were either non-diagnostic or referable to Megalosaurus, with no evidence for a second theropod taxon. The anatomy of these bones cannot be distinguished from M. bucklandii at present. The material from Oakham Quarry is currently under review (Benson 2009a), but some of the bones show autapomorphies of Megalosaurus whereas others (e.g. SDM 44.19, a partial ilium) represent a basal tetanuran of uncertain affinities that is distinct from either Megalosaurus or Cruxicheiros. At least 39 other localities in the British Bajocian–Bathonian have yielded isolated theropod bones and teeth, most of which are indeterminate (Benson 2009a). In summary, three large bodied basal tetanurans, Megalosaurus, Cruxicheiros, and an additional unnamed taxon are known from the early–middle Bathonian of England (Fig. 12). The Callovian Vaches Noires cliffs of northern France have produced several fragmentary theropod specimens in addition to the holotype of Piveteausaurus. Among them is a partial braincase from Auberville (Bülow collection 25192; Knoll et al. 1999) that probably pertains to a megalosauroid, although referral to Allosauroidea cannot be excluded. A pair of articulated frontals from Houlgate (Enos collection; Buffetaut & Enos 1992) can be referred to Tetanurae based on the presence of a distinct socket for articulation with the prefrontal. The anteroposteriorly long dimensions of these frontals suggest placement among megalosauroids or other basal tetanurans rather than avetheropods. Knoll et al. (1999, p. 106) suggested that the specimen might pertain to the same taxon as the braincase, which would further imply that the latter is a megalosauroid. Both elements are very similar to the holotype of Piveteausaurus, especially given our reinterpretation of the sutural contacts in the latter, and we suggest that they may be referable to this form. Finally, a maxilla fragment from Villers-sur-Mer was referred to Megalosaurus sp. (Pennetier collection 380; Buffetaut et al. 1991a). It bears large foramina on the lateral surface and gaps between the roughly surfaced paradental plates that are more consistent with the morphology of megalosauroids than allosauroids, although it cannot be referred to Megalosaurus in particular. There appears to be good evidence for at least two megalosauroids among contemporaneous French and English specimens, a long snouted form (Eustreptospondylus) and

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a shorter snouted form. The second taxon is represented by at least the maxilla, which has a short anterior process, but might also include the holotype of Piveteausaurus (as well as the isolated frontal and braincase). ‘Megalosaurid’ specimens from the Late Oxfordian Sables de Glos of Lisieux, Basse-Normandie, France (Buffetaut et al. 1985) comprise only serrated, laterally compressed teeth that should be regarded as Theropoda indet. Theropod remains are rare in the Kimmeridgian–Tithonian deposits of Europe, known primarily from Portugal (Allosaurus, Aviatyrannis, Ceratosaurus, Torvosaurus) and England (Stokesosaurus and various ‘megalosaur’ remains). Powell (1987, p. 108) referred a worn maxilla fragment from a large theropod that had been dredged from the seabed west of Portland, Dorset (DORCM G10603; Aulacostephanus autissiodorensis Ammonite Zone; lower Kimmeridgian) to Megalosauridae. The presence of an anteroventrally inclined anterior section of dorsal boundary of the paradental plates supports this assignment, and the dorsoventral striations on the paradental plates may allow identification as Megalosaurus (Benson & Barrett 2009). We consider the specimen as Megalosauridae indet., cf. Megalosaurus, and note that it also shares very tall paradental plates with Megalosaurus and Torvosaurus. Fragmentary large theropod remains comprising a right tibia (OUMNH J.29886; Huene, 1926a) and left metatarsus (OUMNH J.13586a–c; Phillips 1871) were also recovered from unknown horizons of the Kimmeridge Clay Formation (Kimmeridgian–early Tithonian) at Swindon, Wiltshire. The half-century interval between these reports suggests that they were not found in association. All were referred to Megalosaurus bucklandii (Phillips 1871; Huene 1926a; Delair 1973), although Huene (1932) later revised this to Megalosaurus (?) sp. The proportions of the tibia are similar to those of Torvosaurus, the most robust basal tetanuran, but there is little else to allow firm identification and they are regarded here as Tetanurae indet. These specimens are consistent with the presence of a robust megalosaurid in the UK, contemporaneous (and possibly conspecific) with the Portuguese Torvosaurus. Many other theropod elements from the Kimmeridge Clay Formation should be regarded as Theropoda indet., including phalanges from the lower Kimmeridgian Rasenia cymodoce Zone of Wyke Regis, Dorset (Brokenshire & Clarke 1993; Benson & Barrett 2009), a tooth from Foxhangers, Wiltshire referred to Megalosaurus insignis (NHMUK 46388; Lydekker 1888), ‘megalosaur’ phalanges from an unspecified locality in Wiltshire (DZSWS 3009; Delair 1973), and a proximal caudal vertebra from Shotover, Oxfordshire (OUMNH J.47134; Powell 1987). The fragmentary pectoral remains of a large theropod from the early Valanginian of Montmirat, France were

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ENGLAND

FRANCE

fittoni

Tithonian

rotunda pallasiodes pectinatus

Stokesosaurus

hudlestoni wheatleyensis Compsognathus

scitulus

Kimmeridgian

autissiodorensis

megalosaurid

eudoxus

megalosaurid, tetanuran

mutabilis cymodoce baylei pseudocordata

Oxfordian

decipiens cantisnigrae

Eustreptospondylus

plicatilis cordatum mariae

Metriacanthosaurus Streptospondylus

Bathonian

Callovian

lamberti

Bajocian


elegans

athleta

Eustreptospondylus

Piveteausaurus, megalosaurids

coronatum jason calloviense macrocephalus discus aspidoides hodsoni morrisi subcontractus progracilis tenuiplicatus zigzag parkinsoni garantiana subfurcatum humphriesianum sauzei laeviuscula discites

tetanuran Proceratosaurus Megalosaurus, Iliosuchus

Dubreuillosaurus, Poekilopleuron

Megalosaurus, Cruxicheiros, tetanuran Duriavenator Magnosaurus

Figure 12. Stratigraphical relationships of Jurassic tetanurans in England and France. The central columns show the approximate stratigraphic position of known theropod localities (small circles). Where possible, individual taxa have been identified to the left (England) and right (France) of these columns. Symbols: triangles, megalosauroids; squares, allosauroids and other non-coelurosaurian avetheropods; stars, coelurosaurs.


The phylogeny of Tetanurae described by Pérez-Moreno et al. (1994) and interpreted as a close relative of Allosaurus. The specimen, which included a partial scapula, humerus and manus, displays proportions that are similar to those of Allosaurus but also other basal tetanurans (e.g. Afrovenator). Three characters were used to link it with Allosaurus (humerus/scapula length ratio 1 tonne) theropods are still the early Late Jurassic metriacanthosaurids, but the largest allosaurids are approximately coeval with giant megalosaurs in the upper levels of the Morrison Formation (Foster 2003), as well as the large (unidentified) theropods of the Tendaguru. Giant carcharodontosaurians (including ‘acrocanthosaurs’) occur from Aptian through at least Coniacian times, contemporaries of the giant terrestrial spinosaurids. The largest tyrannosaurids do not occur until the Campanian–Maastrichtian, when the largest abelisaurids are also known from South America. Thus it was quite possible for more than one giant theropod to coexist, even in the same palaeoenvironment, but this may have been dependent on significant variance in feeding habit between taxa. Finally, although understanding the evolution of large size has greater scientific value than the absolute sizes of theropods per se, it is the latter that has garnered almost the entirety of popular and scientific attention since the original discovery of Tyrannosaurus rex. The emphasis on absolute sizes has had the unfortunate effect of sidelining many questions of genuine biological interest, including the pattern and process of size increases, their overall magnitude (i.e. from ancestor to descendant), apparent repetition, and possible upper limits. As an exception, Bakker et al. (1992) focused on how each large theropod family independently reached nearly the same maximum size. This remains true even in light of the many new discoveries of giant theropod remains made since. Indeed, the difference in size between the skulls of the largest carcharodontosaurids and tyrannosaurids amounts to only a few centimetres, well within the range of species variation and rendering scientifically moot the question of which is the ‘largest theropod’. Interestingly, these taxa exhibited different overall body dimensions, fore- and hind limb morphologies, and tooth shape, which implies quite different predatory habits. Body mass (rather than

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M. T. Carrano et al. Jurassic Early

He

Si

Pl

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To

Aa

Bj

Late

Bt Cl

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Ki

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Ca

Ma sampling

‘D.’ sinensis Cryolophosaurus

1

Chuandongocoelurus Monolophosaurus Marshosaurus Condorraptor Piatnitzkysaurus Eustreptospondylus Streptospondylus Duriavenator Megalosaurus Torvosaurus Leshansaurus Afrovenator

2

Hidden Lake


Magnosaurus

6

4

Piveteausaurus Dubreuillosaurus Poekilopleuron 3 Xuanhanosaurus Shidaisaurus Y. zigongensis

Baryonyx Suchomimus Angaturama Irritator Spinosaurus Y. shangyouensis S. hepingensis

S.dongi Metriacanthosaurus Siamotyrannus 7

Saurophaganax Allosaurus

Erectopus Neovenator Chilantaisaurus

8 Megaraptor Aerosteon Fukuiraptor

Orkoraptor Australovenator

5 Concavenator 9

Eocarcharia Acrocanthosaurus Shaochilong Tyrannotitan Carcharodontosaurus Giganotosaurus Mapusaurus

COELUROSAURIA

Figure 13. Stratigraphically calibrated phylogeny, based on current results. Note that coelurosaurs and taxa outside Tetanurae have been omitted for clarity. Solid bars represent range estimates (including uncertainty) for taxa included in the present study; open bars represent range extensions based on taxa and specimens assigned to the respective clades; solid lines represent missing lineages inferred from the topology; dashed lines represent tentative assignments. Filled circles along time line represent known theropod temporal occurrences, based on information from the Paleobiology Database (www.paleodb.org). Clade names in circles: 1, Piatnitzkysauridae; 2, Megalosauridae; 3, Spinosauridae; 4, Tetanurae; 5, Avetheropoda; 6, Metriacanthosauridae; 7, Allosauridae; 8, Neovenatoridae; 9, Carcharodontosauridae.

‘size’) also varies, with the largest Tyrannosaurus rex specimens exhibiting limb proportions that suggest substantially heavier masses than carcharodontosaurids of similar linear dimension. Bipedalism may have constrained the maximum size attainable by theropods (Bakker et al. 1992; Farlow et al. 1995), but this hypothesis requires further testing. Diversity. At a minimum, tetanurans originated early in the Early Jurassic (Fig. 13), as evidenced by ‘Dilophosaurus’ sinensis and Cryolophosaurus, as well as the sister taxon of Tetanurae, Ceratosauria, which is first represented by Berberosaurus (Pliensbachian–Toarcian; Allain et al. 2007). Given the presence of coelophysoids in the Late Triassic, this implies the existence of stem-averostrans in the Carnian. Of course, as this is a minimum divergence estimate it is also possible that both the ceratosaur and teta-

nuran lineages existed even as early as the Late Triassic. With the currently known samples of these clades, the 95% confidence interval on the first occurrence of Tetanurae (but not Ceratosauria) extends into the Norian (using the method of Marshall 1994, which does not assume random distribution of samples) (Fig. 14). Regardless of the exact time of origin, the earliest tetanurans diversified alongside two other distinct theropod clades, coelophysoids and ceratosaurs. These three clades coexisted through the Early Jurassic, but coelophysoids apparently went extinct at the close of this interval or shortly thereafter (Carrano & Sampson 2004). Neither ceratosaurs nor tetanurans appear to have been particularly diverse during the Early Jurassic, but the poor quality of the fossil record from this interval, especially in terms of geographical coverage, suggests that this may be partly artefactual.

The phylogeny of Tetanurae Jurassic Early He

Si

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Cretaceous Middle

To

Aa

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Bj

Bt Cl

Late Ox

Ki

Early Ti

Be Va

Ha

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Late Ap

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Co Sa

Ca

Ma

‘D.’ sinensis Cryolophosaurus Chuandongocoelurus Monolophosaurus Piatnitzkysauridae Megalosauridae

Hidden Lake Spinosauridae Metriacanthosauridae

Allosauridae

Erectopus


Neovenatoridae Carcharodontosauridae COELUROSAURIA

Figure 14. Stratigraphically calibrated phylogeny showing sampling density for major clades, simplified from Fig. 13. Black circles represent individual occurrences; white circles represent tentative assignments; black bars represent densely sampled intervals; white bars extending beyond noted samples represent range uncertainties.

The minimum divergence times of subsequent tetanuran clades suggest a radiation no later than the late Early Jurassic (Fig. 13). Subsequent to their initial appearance, Tetanurae radiated into two main clades, Megalosauroidea and Avetheropoda. The earliest megalosauroids are known from the Bajocian (Duriavenator, Magnosaurus), implying the presence of basal avetheropods at this time also. Definitive avetheropod fossils occur slightly later in the Middle Jurassic of Europe (Bathonian: Proceratosaurus). In the Middle Jurassic of China, ‘stem’ tetanurans (Chuandongocoelurus, Monolophosaurus) persisted alongside megalosaurids (Leshansaurus) and metriacanthosaurid allosauroids, the first occurrences of which (Shidaisaurus) are penecontemporaneous with those of avetheropods in Europe. By the Callovian and Late Jurassic, the fossil record demonstrates the widespread presence of multiple clades within both megalosauroids and avetheropods. Among the former are megalosaurids and piatnitzkysaurids, implying the presence of spinosaurids as well. Indeed, Eustreptospondylus and Streptospondylus are ambiguously resolved as basal megalosaurians and may therefore represent the most basal spinosaurids (although they could equally well be megalosaurids). Avetheropodans included allosaurids and metriacanthosaurids, therefore implying the presence of basal carcharodontosaurians, which may be represented in the Tendaguru Formation of Tanzania. The earliest definite spinosaurid fossils are possibly Late Jurassic (Tendaguru Formation) but certainly mid-Neocomian in age; the earliest definite carcharodontosaurian fossils

are possibly also Late Jurassic (Tendaguru Formation), but certainly Barremian (Concavenator, Neovenator). Fossils of both groups are abundant in the ‘medial’ Cretaceous interval. Allosaurids are an unexpectedly small group, restricted to Allosaurus and Saurophaganax and present with certainty only in the Kimmeridgian–Tithonian of North America and Europe. Metriacanthosaurids are almost exclusively known from the Oxfordian of Asia and Europe. It is not yet possible to determine whether metriacanthosaurids were present in the Oxfordian of North America but replaced by allosaurids in the Kimmeridgian, as seems to have occurred in Europe. Alternatively, the distributions of these two clades may reflect genuine biogeographical differentiation. Among the megalosauroids, only spinosaurids definitively survived the Early Cretaceous, although there may have been a relictual megalosauroid in Late Cretaceous Antarctica (Hidden Lake specimen). Afrovenator was the sole non-spinosaurid megalosauroid known for this interval, but recent work suggests that it may be considerably older (Rauhut & López-Arbarello 2009). Although their initial diversification included nearly all the continents, megalosauroids may have been restricted to Europe and Gondwana by the late Early Cretaceous. The paucity of spinosaurid discoveries from the late Early and early Late Cretaceous may reflect genuinely sparse and geographically patchy fossil sampling. Allosauroids were also geographically widespread in their early history, and they too are primarily known from Gondwanan landmasses by the Late Cretaceous. Chilantaisaurus and Shaochilong



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Figure 15. Global distributions of non-coelurosaurian tetanuran theropods. A, Early Jurassic (map reconstruction showing continents at ca 200 Ma); B, Middle Jurassic (170 Ma); C, Late Jurassic (150 Ma); D, earliest Cretaceous (120 Ma); E, medial Cretaceous (90 Ma); F, latest Cretaceous (65 Ma). Symbols: circles, non-tetanuran, coelurosaur and indeterminate theropods; bulls-eyes, basal averostrans and questionable or indeterminate basal tetanurans; squares, megalosauroids; triangles, allosauroids. Maps copyright Ron Blakey, Northern Arizona University.



Figure 15. (Continued)

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Figure 16. Stratigraphical fit of theropod taxa in this analysis, using age rank and clade rank. Open circles, basal theropods; grey circles, megalosauroids; filled circles, avetheropods.

are known from the ?Turonian of China, but the record of pre-Campanian Late Cretaceous theropods from Laurasia is otherwise very poor. It is likely that the Laurasian radiation of large bodied coelurosaurs, known primarily from the Campanian and Maastrichtian, came at the eventual expense of more basal tetanuran lineages. South American megaraptoran allosauroids are the only Campanian–Maastrichtian basal tetanurans. Finally, it is interesting to note how common it is to find multiple lineages of apparently contemporaneous large theropods within single formations of Jurassic and Early Cretaceous age. Although the Morrison Formation of western North America has been noted for its unusually high diversity of large theropods (at least five; Chure et al. 2000), other locales have produced at least three contemporaneous taxa. In addition to the Late Jurassic beds of Tanzania and Portugal, these include the Bathonian (a megalosaurid and basal tetanurans) and Berriasian (allosauroids, a spinosaurid) of the United Kingdom, the Albian and Cenomanian of North Africa (spinosaurids, carcharodontosaurids, ceratosaurs), the Allen Formation of Argentina (basal tetanurans, carcharodontosaurids, abelisaurids), and perhaps the Adamantina Formation of Brazil (?spinosaurids, ?carcharodontosaurids, abelisaurids). Many additional formations document the presence of at least two large theropods, but most of these are far from exhaustively sampled. Biogeography. The biogeographical history of various theropod groups has received intense attention in recent years (e.g. Carrano et al. 2002; Sereno et al. 2004), but little consensus has been achieved. It is beyond the scope of this paper to review the major hypotheses and analyses, even briefly. Unlike phylogenetic analyses, there are significant discrepancies in terminology, data sampling, and

methodology that make comparisons between results problematic. Instead, we use the results of the present phylogenetic analysis to suggest a substantial revision to the null models currently employed by many biogeographical studies. As noted in a previous paper (Benson et al. 2010), most theropod clades achieved a broad geographical distribution quite early in their geological history. It is this early history that should inform considerations of what the neutral expectations should be for any biogeographical scenario. With this in mind, we suggest that any diverse clade appearing prior to the Late Jurassic should be suspected of having a global distribution a priori, at least on the continental geographical scale. Given the incompleteness of the fossil record, it is therefore especially important to determine genuine absences, because well-supported evidence of absences will be particularly enlightening. For example, the absences of sauropods and large non-tyrannosaurid theropods from the Hell Creek Formation of North America appear to be genuine, based as they are on the lack of fossils of these forms among the hundreds of thousands collected thus far. The presence of both groups in earlier strata, and their survival elsewhere in the Maastrichtian, indicates that a regional extinction is responsible for this pattern. Regional extinctions must be more important than has been suspected in determining the later Mesozoic patterns of dinosaur distribution. As discovery and analysis have advanced in recent decades, most known clades have been identified in progressively older strata, and indeed nearly all major dinosaur clades (e.g. Ceratopsia) are now known from pre-Late Jurassic rocks. Theropods conform to this trend, with definite coelurosaurs (and therefore all more primitive theropod clades) now well identified from the Middle Jurassic, and several clades that probably originated in the Jurassic (metriacanthosaurids, megalosauroids, carcharodontosaurians) showing an early widespread distribution (Fig. 15). In this respect, the modern world and the distributions of its terrestrial taxa constitute a poor model for Mesozoic continental biogeography, because most extant clades had their origins in a world of fragmented continents. Even so, many modern clades whose origins are known to predate significant continental breakup (e.g. Marsupialia/Metatheria, Pleurodira, Anguimorpha) have fossil records that are nearly global in breadth. We suggest a model wherein the initial supposition is for large Mesozoic clades to have had the potential to achieve a cosmopolitan distribution, which would befit an early Mesozoic world in which continental connection was the rule. This operates under the assumption that organisms quickly reach their maximum potential dispersion, at least on geological time frames, and sets the groundwork for interpreting observed taxon distributions. It does not imply


The phylogeny of Tetanurae that all dinosaur clades had global distributions; rather it is a first-order expectation that provides a framework against which deviations can be identified and studied. Second, it is critical to assess the quality of the fossil record. Given recent advances in understanding the dinosaur fossil record and the steady rate of discovery of new taxa, it is not tenable merely to assume that all taxon absences are real in every locale. The tendency to do so involves the tacit assumption that the fossil record is mostly complete and/or accurate, and therefore does not require as rigorous analytical treatment as phylogenetic hypotheses. (One would not support coding every ‘?’ as ‘0’ in a phylogenetic analysis, but this is analogous to assuming all taxon absences are genuine in the fossil record.) With this information in hand, it becomes possible to examine biogeographical patterns. Deviations from prior expectations can be analysed in more detail in the light of the known fossil record. Only at this stage is it appropriate to introduce second-order geographical barriers (e.g. mountain ranges, deserts, or other features that impose differential restrictions on different terrestrial organisms). We know next to nothing about the actual dispersal potentials for any dinosaurs with respect to barriers of this kind, and so they cannot be invoked as independent causative factors unless it can first be demonstrated that taxon absences are likely to be real. This is not to say that such barriers were not important in creating biogeographical patterns amongst dinosaurs in the Mesozoic – certainly they were. But our knowledge of them, and of how dinosaurs interacted with them, is insufficient to permit them a role in setting prior expectations. Stratigraphical fit. As with other dinosaur clades, there is relatively good overall fit between the sequence of clade appearances and their stratigraphical first appearance among tetanurans. This is reflected in the highly significant correlation of age rank and clade rank based on our phylogenetic hypothesis (rho = 0.73, p < 0.001), as well as the graphical comparison of these two variables (Fig. 16) and the SCI of 0.605. The RCI is quite low (70.77%), emphasizing how much of the predicted record of tetanurans remains to be discovered. Few major discrepancies occur in sequence; rather there are extensive missing lineages throughout the phylogeny. Significant among these are the lineages leading to Spinosauridae (∼ 42 Ma missing), Allosauria (∼ 20 Ma), Spinosaurinae (∼ 20 Ma), and Carcharodontosauria (∼ 23 Ma). The sampling gaps in late Early–early Middle and earliest Cretaceous times are apparent in the absence of taxa with appropriate age ranks (7–8 and 15–16, respectively). The late appearing taxa with low clade rank are derived ceratosaurs, whose ranks are artificially low due to the low numbers of taxa sampled for this study. In addition, the two major ‘pulses’ of tetanuran evolution (whether genuine or artefactual) are evident as discrete clusters of taxa in the centre and upper-right portions of

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the graph. These reflect the Jurassic radiation of megalosauroids and the Cretaceous radiations of spinosaurids and derived allosaurians.

Conclusions Tetanuran theropods represent the majority of predatory dinosaurs, but their early evolutionary history has remained difficult to decipher. Although many taxa are known from fragmentary remains, several recent discoveries combined with a re-examination of existing materials has allowed us to significantly improve phylogenetic resolution for this important group. Here we present the results of a detailed, species-level phylogenetic analysis of 59 ingroup taxa, mostly tetanurans. Our study utilizes 351 characters derived from firsthand study of specimens and comprehensive review of all prior systematic analyses. The results support the successive placement of Ceratosauria and Tetanurae as more derived groups relative to Coelophysoidea. Several taxa are found to occupy ‘stem’ positions relative to Tetanurae, including two (Cryolophosaurus and ‘Dilophosaurus’ sinensis) that were previously considered to be more basal. Within Tetanurae, Megalosauroidea and Avetheropoda are sister taxa. The former clade is quite diverse, with two Jurassic clades (Piatnitzkysauridae fam. nov. and Megalosauridae) and the Cretaceous Spinosauridae. Within Megalosauridae, our results support the presence of two subfamilies, Megalosaurinae and Afrovenatorinae subfam. nov. Avetheropoda includes the Jurassic Metriacanthosauridae and Allosauridae as well as the Cretaceous Carcharodontosauria (comprising Neovenatoridae and Carcharodontosauridae). The presence of Early and Middle Jurassic forms on the tetanuran ‘stem’ lineage requires a minimum appearance time of Late Triassic or earliest Jurassic for Averostra, and the early Middle Jurassic appearance of several true tetanurans (Duriavenator, Magnosaurus) suggests that the radiation of basal megalosauroids and avetheropods took place not later than the late Early Jurassic. Given the relatively poor Early Jurassic record, it is certainly possible that these lineages extend considerably farther back in time. Substantial missing lineages are present for Spinosauridae and Allosauria, although the > 20 million-year gap for Carcharodontosauria may be mitigated by the presence of a Tendaguru form. In addition, we review all previously published tetanuran taxa and occurrences in order to document thoroughly and assess the fossil record of the group. Many of these taxa were too fragmentary to be included in our analysis, but can nonetheless be identified as belonging to Tetanurae or particular ingroups. Although some records of tetanurans can be refuted, the remaining sample documents a much broader geographical and temporal radiation than the


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phylogenetic results alone. A revised classification of tetanurans is given in the Appendix. Tetanuran evolution seems to exhibit a series of ‘waves’ of diversification, but it is difficult to determine whether these are relatively discrete pulses or an artefact of uneven sampling. Regardless, the dominance of particular groups, particularly at large body sizes, follows a loose succession: Megalosauridae, Piatnitzkysauridae and Metriacanthosauridae (Middle Jurassic); Allosauridae, Megalosauridae and Metriacanthosauridae (Late Jurassic); Spinosauridae, Carcharodontosauridae and Neovenatoridae (Early and mid Cretaceous). Terminal Cretaceous terrestrial ecosystems appear to have been dominated by a combination of large-bodied tetanurans (i.e. tyrannosaurid coelurosaurians; primarily Northern Hemisphere) and abelisaurid ceratosaurians (primarily Southern Hemisphere). The morphological evolution of tetanurans is complex, as would be expected of a radiation involving several contemporaneous clades. General skull proportions vary between a more primitive design with greater skull roof elaboration (common in allosauroids and outside Tetanurae) and a more elongate form that lacks these developments (more typical among megalosauroids). Spinosaurids show the only significant departures in tooth morphology, snout configuration, and morphology of the posterior skull within noncoelurosaurian tetanurans. Although some basal tetanurans (e.g. Neovenatoridae) display parallels with coelurosaurs, in general the group is characterized by a relatively conservative limb morphology that shows little tendency towards the extremes of locomotor evolution. Tetanuran evolution records a gradual transition from a primitive theropod condition to the more bird-like features common among coelurosaurs, although a number of ‘avian’ features were already present in many tetanuran taxa. Basal tetanurans achieved giant body sizes in a number of lineages, reaching very similar maximum sizes despite this independent evolution. The presence of multiple large theropods in several different palaeoenvironments suggests some degree of ecological resource partitioning. The biogeographical history of tetanurans is likewise complex, spanning more than 110 Ma and all continents. At present the density of sampling is insufficient, both temporally and geographically, to permit detailed analysis of tetanuran biogeographic evolution. However, we suggest an alternative model for assessing the history of the group, and of terrestrial vertebrates more generally during this time interval. The presence of all major lineages of tetanurans prior to any significant breakup of Pangaea implies that these clades would have had the opportunity to disperse widely and achieve cosmopolitan distributions. Their absences from regions later in time would then be due to regional extinctions or dispersal failure. This model should be appropriate for any early Mesozoic radiation;

a dispersal/vicariance dichotomy is an appropriate ‘null model’ only for more highly nested groups whose origins postdate continental fragmentation.

Acknowledgements We would like to thank the many people who allowed us access to specimens in their care, including: Peter Wellnhofer (BSP); Rod Scheetz and Brooks Britt (BYU); Li Kui (CCG); Amy Henrici (CMNH); Kenneth Carpenter (DMNH); Dan Chure (DNM); William Simpson (FMNH); Wolf-Dieter Heinrich and David Unwin (MB); Zhao Xijin and Xu Xing (IVPP); John Foster (MWC); José Bonaparte, Alejandro Kramarz and Fernando Novas (MACN); Rodolfo Coria (MCF); Leonardo Salgado (MC); Diego Pol (MCF); Sean Duran (Miami Science Museum); Steve Hutt and Martin Munt (MIWG); Octavio Mateus (ML); Philippe Taquet, Daniel Goujet, Ronan Allain and Emily Long (MNHN); Rodolfo Coria (MPEF); Jorge Calvo (MUCPv); Juan Canale (MUCPv-Ch); Vince Schneider and Drew Eddy (NCSM); Jeff Person (OMNH); Paul Jeffery (OUMNH); Jaime Powell (PVL); Oscar Alcober and Ricardo Mart´ınez (PVSJ); Jaime Powell (PVL); David Mullin (SDM); Rich Ketcham and Tim Rowe (TMM); Sandra Chapman, Angela Milner and Paul Barrett (NHM); Robert Masek and Paul Sereno (University of Chicago); Pat Holroyd and Kevin Padian (UCMP); Mike Getty (UMNH); and Guang-Zhou Peng (ZDM). In several cases, we were granted access to then-unpublished materials, for which we are particularly grateful. This paper benefited from numerous discussions with Jeffrey Wilson, Cathy Forster, Ron Tykoski, Paul Barrett, David Norman, and Nathan Smith, careful editorial work by Susannah Maidment, and detailed and insightful reviews by Oliver Rauhut and Stephen Brusatte. Translations of Allain (2001), Bonaparte & Novas (1985), Bonaparte (1986), Depéret & Savornin (1925, 1928), Dong et al. (1983), Eudes-Deslongchamps (1837), Janensch (1925), Kurzanov (1989), Lapparent (1960), Nessov (1995), Novas (1991a,b, 1992), Powell (1979), Stromer (1915, 1931), Taquet & Welles (1977) and Thévenin (1907) are available at the Polyglot Paleontologist website (http://www.paleoglot.org/). This research was supported by NSF DEB-9904045 to MTC and SDS; NERC studentship NER/S/A/2005/13488 to RBJB; and smaller grants from The Jurassic Foundation, SYNTHESYS and the Palaeontographical Society to RBJB.

Supplementary material Supplementary material is available 10.1080/14772019.2011.630927

online

DOI:



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291

Appendix 1


Taxonomic hierarchy of Theropoda derived from the results of the present analyses. For each family-level taxon, the original author(s) of the first associated family-level name is given immediately after the name, followed by the author(s) who first created the name at that specific rank (if different). For example, Coelophysinae was named as a subfamily by Nopcsa (1928) but first elevated to family rank by Paul (1988a), so we list Coelophysidae (Nopcsa, 1928) Paul, 1988a. NEOTHEROPODA Bakker, 1986 Coelophysoidea Holtz, 1994 Dilophosaurus wetherilli Welles, 1970 Coelophysidae (Nopcsa, 1928) Paul, 1988 Coelophysis Cope, 1889 C. bauri (Cope, 1887a) Cope, 1889 C. rhodesiensis (Raath, 1969) Paul, 1988a AVEROSTRA Paul, 2002 (sensu Ezcurra 2006) CERATOSAURIA Marsh, 1884 Elaphrosaurus bambergi Janensch, 1920 Ceratosaurus nasicornis Marsh, 1884 Abelisauroidea (Bonaparte & Novas, 1985) Bonaparte, 1991 Masiakasaurus knopfleri Sampson, Carrano & Forster, 2001 Majungasaurus crenatissimus (Depéret, 1896) Lavocat, 1955 TETANURAE Gauthier, 1986 [syn. Avipoda Novas, 1992] ‘Dilophosaurus’ sinensis Hu, 1993 Cryolophosaurus ellioti Hammer & Hickerson, 1994 Monolophosaurus jiangi Zhao & Currie, 1994 Chuandongocoelurus primitivus He, 1984 ORIONIDES nom. nov. Megalosauroidea (Fitzinger, 1843) Walker, 1964 [syn. Torvosauroidea (Jensen, 1985) Sereno et al., 1994; Spinosauroidea (Stromer, 1915) Olshevsky, 1995] Piatnitzkysauridae fam. nov. Condorraptor currumili Rauhut, 2005b Marshosaurus bicentesimus Madsen, 1976b Piatnitzkysaurus floresi Bonaparte, 1979 Megalosauria Bonaparte, 1850 Streptospondylus altdorfensis Meyer, 1832 (Megalosauria incertae sedis) Spinosauridae Stromer, 1915 Baryonychinae (Charig & Milner, 1986) Sereno et al., 1998 Baryonyx walkeri Charig & Milner, 1986 Suchomimus tenerensis Sereno et al., 1998 Spinosaurinae (Stromer, 1915) Sereno et al., 1998 Angaturama limai Kellner & Campos, 1996 Irritator challengeri Martill et al., 1995 Spinosaurus aegyptiacus Stromer, 1915 Megalosauridae Fitzinger, 1843 Eustreptospondylus oxoniensis Walker, 1964 Afrovenatorinae subfam. nov. Afrovenator abakensis Sereno et al., 1994 Dubreuillosaurus valesdunensis (Allain, 2002) Allain, 2005a Leshansaurus qianweiensis Li et al., 2009 Magnosaurus nethercombensis (Huene, 1923) Huene, 1932 Piveteausaurus divesensis (Walker, 1964) Taquet & Welles, 1977 Poekilopleuron bucklandii Eudes-Deslongchamps, 1837 Megalosaurinae (Fitzinger, 1843) Paul, 1988 Duriavenator hesperis (Waldman, 1974) Benson, 2008


292


Megalosaurus bucklandii Mantell, 1827 Torvosaurus tanneri Galton & Jensen, 1979 AVETHEROPODA Paul, 1988 [syn. Neotetanurae Sereno et al., 1994] Allosauroidea (Marsh, 1878) Currie & Zhao, 1994 Metriacanthosauridae Paul, 1988a [syn. Sinraptoridae Currie & Zhao, 1994] Xuanhanosaurus qilixiaensis Dong, 1984 (Metriacanthosauridae incertae sedis) Yangchuanosaurus Dong, Chang, Li, & Zhou, 1978 Y. shangyouensis Dong, Chang, Li, & Zhou, 1978 Y. zigongensis (Gao, 1993) n. comb. Metriacanthosaurinae (Paul, 1988a) subfam. nov. Metriacanthosaurus walkeri (Huene, 1923) Walker, 1964 Shidaisaurus jinae Wu et al., 2009 Siamotyrannus isanensis Buffetaut, Suteethorn & Tong, 1996 Sinraptor Currie & Zhao, 1994 S. hepingensis (Gao, 1992) Currie & Zhao, 1994 S. dongi Currie & Zhao, 1994 Allosauria Paul, 1988 Allosauridae Marsh, 1878 [syn. Labrosauridae Marsh, 1882; Antrodemidae Stromer, 1934] Allosaurus Marsh, 1877 A. europaeus Mateus et al., 2006 A. fragilis Marsh, 1877 A. jimmadseni Chure et al., 2006 Saurophaganax maximus Chure, 1995 Carcharodontosauria Benson et al., 2009 Neovenatoridae Benson et al., 2009 Neovenator salerii Hutt, Martill & Barker, 1996 Chilantaisaurus tashuikouensis Hu, 1964 Megaraptora Benson et al., 2009 Aerosteon riocoloradensis Sereno et al., 2008 Australovenator wintonensis Hocknull et al., 2009 Fukuiraptor kitadaniensis Azuma & Currie, 2000 Megaraptor namunhuaiquii Novas, 1998 Carcharodontosauridae Stromer, 1931 Acrocanthosaurus atokensis Stovall & Langston, 1950 Concavenator corcovatus Ortega, Escaso & Sanz, 2010 Eocarcharia dinops Sereno & Brusatte, 2008 Shaochilong maortuensis Brusatte et al., 2009 Carcharodontosaurinae (Stromer, 1931) subfam. nov. Carcharodontosaurus Stromer, 1931 C. iguidensis Brusatte & Sereno, 2007 C. saharicus (Depéret & Savornin, 1925) Stromer 1931 Giganotosaurus carolinii Coria & Salgado, 1995 Mapusaurus roseae Coria & Currie, 2006 Tyrannotitan chubutensis Novas et al., 2005 COELUROSAURIA Huene 1914 Lourinhanosaurus antunesi Mateus, 1998 Compsognathus longipes Wagner, 1861 Ornitholestes hermanni Osborn, 1903 Proceratosaurus bradleyi (Woodward, 1910) Huene, 1926


293

Appendix 2 Taxon-by-character matrix for the 61 taxa used in this study. The two outgroup taxa are listed first, followed by the 59 ingroup taxa listed alphabetically. Characters are scored from 0 to 1, 2, 3, or 4. Higher numbers represent uncodable character observations for particular taxa. Polymorphisms were assumed to represent uncertainties.


Eoraptor 00010 000?0 0000? ????? 01010 00?0? 00100 00?00 00100 ??000 1?000 00010 00000 ?0000 00000 0?10? 00??0 ?0010 00?00 00?0? ????? 0??00 ?0?10 10000 00?0? 0000? 00?00 00?0? 00?00 01000 00000 00090 ????? ????8 00000 00000 0000? 0?000 00000 00000 0?000 0001? 00100 00?00 00010 00000 10100 00000 00?00 10000 01000 00000 00001 00000 00?00 00000 0000? ?000? 00000 00?00 000?0 000?0 00?00 00000 00??? 80000 00000 00000 00000 00000 0 Herrerasaurus 00001 00000 0000? ???0? 00010 00?0? 0?000 00?00 00100 ??001 20100 00000 00000 ?0000 01000 00000 020?0 00010 ?0000 0?10? ?0000 0000? 00000 0?100 00?0? 0000? 00001 00100 00?0? 01000 00000 00080 0?000 010?9 00000 00000 0000? 0?100 0000? 00000 00000 000?? 001?? 00?1? 011?0 ??000 10100 00001 00000 10100 01000 00000 00001 00000 00?00 00000 0000? 0?001 00000 0?000 00??0 00010 00000 00000 001?? 90100 00100 00000 00000 00000 0 Acrocanthosaurus 020?1 100?0 00000 11110 00000 21111 11000 10?00 00002 11101 21110 00202 12011 ?1121 11102 0110? 02??1 10121 00000 10011 01000 12011 111?1 11002 10010 10200 10?11 111?0 00100 01000 00100 11012 ?0011 0011? 00?00 12110 00100 01100 11211 110?0 02100 00001 01?1? 11011 01200 12001 210?0 01011 11000 01131 10111 11100 0???? 010?1 0??11 10110 20120 1?120 ?2011 11000 22011 10111 10000 11001 0312? 1010? 1110? 1??01 21011 1001? 1 Aerosteon ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?01?0 00?11 1???? ????? ?1000 0200? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 11012 1???? ???11 ?0100 1?1?0 00100 01011 01011 1???? ?2?11 0??0? ????? ???01 01100 121?? ????? ????? ????? ????? ????? ????? 01011 01001 02211 10110 20120 1?120 ????? ????? ????? ????? ????? ?1001 ?412? ???01 11202 1110? ????? ????? ? Afrovenator 01011 000?? ????? ????? 001?0 00001 00111 1?100 01000 021?1

????? ????? ????? ????? 001?1 10101

????0 ??00? ??000 ??100 0?011 10???

01010 00111 00000 000?? 01011 11101

11000 1001? 01??? ?1?01 ?011? 1?011

00100 ????? ??10? 010?? 2012? 100??

12001 ????? 11001 ????? 1?00? 1

1000? ????? ?0?01 ???01 00101

????0 ????? 10000 21??? 11000

?0002 ????? 10001 ??00? 11101

01001 ????? 10110 0???? 10000

?0110 ????? 0001? ?0?31 11000

Allosaurus 00101 [01]0000 01000 10100 00000 21111 11000 11011 00002 [01]2[01]00 10110 00000 01011 11100 01102 00000 02001 10110 00110 10010 01020 12011 11111 11200 01001 10201 11111 11100 00100 01102 00100 11001 10011 0010[01] 10000 10110 00100 00100 01011 11000 011?0 00000 0101? 11011 01200 12001 21010 11000 01000 11131 00111 11100 00111 01001 02111 10110 20120 1?110 02001 11000 [12]1011 10111 10000 11001 02122 10101 11102 11101 21011 10010 1 Angaturama 11??? 201?1 121?1 1???0 ????? ????? ???0? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????1 ?0??? ????? ????? ????? 11011 17013 11??? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ? Australovenator ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????

294 ?0000 ????? 01?11 11001

M. T. Carrano et al. 1???? ????? 11??1 14122

????? ????? 01??? 10101

??100 ????? ????? 11212

00100 ????0 ????? 11101

0110? ????? ????? 211??

????? ????? ????? ??010

????? ????? ????? ????? ????? ????? ????? ????? ????? ???00 01010 1???? ????? ????? ????? 22012 10111 10001 ?


Baryonyx 111?0 20101 12101 10?00 00??? 02??? 1??01 ????0 ?0012 02011 ?0??? ???0? ?0?11 11100 ????? ?0?11 01101 00010 11?01 ??010 210?0 11??? ????? ??001 10101 0?[12]10 1???1 ??010 01011 00013 00101 1100[12] ?1001 00100 10001 10111 11010 00110 00101 010?0 ???00 0?00? ?1??1 ?00?? 0?1?0 10001 21111 00010 01101 0???? ????? ???10 ?0?11 010?1 01?11 10110 [01]0?2? 1?0?? ??001 1000? ???1? ???00 11010 0???? ????1 10?0? ????? ????? ??0?? ????? ? Carcharodontosaurus ????? ?0??0 00000 11110 00010 01101 11010 11110 00?02 ?1?0? ?1111 00??2 12112 11?21 1???? 0???? ????1 10121 ?0??0 100?? 01121 121?? ?1??? ????2 ?10?? ????? ????? ??100 00200 00??? ??10? 110?2 ????? 0??01 ?0?00 101?0 ???1? ???0? ????? ????? ???10 0?0?? ?1??? 1???? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?1??1 ????? ??11? ????0 ????? 0??11 ????? 22??1 10111 1??01 ?100? ????? ?010? ????? ????? ????? ????? ? Ceratosaurus 02101 00010 00000 00000 00000 01010 00101 20?00 00002 02001 10000 00010 00011 11110 00101 00100 10001 01010 00000 00110 10000 11100 00?10 00100 00001 ??100 00001 00100 01000 00101 00100 01102 1?010 00100 00000 02110 00000 10100 00101 0211? ?1000 0210? 01000 01101 01111 01001 00110 000?? ??0?? 1??1? 01?00 010?0 10?11 100?0 00011 10111 10110 10000 1[01]000 10011 00110 [01]0011 02100 01010 02002 11101 00102 1??1? 0001? ?10?? ? Chilantaisaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ???01 20110 000?? ????? ????? ????? ???11 ????? ?10?? ??211 0???? ????? ????? ????? ????? 22??1 11??? 1000? 1100? ?4122 1???? 1???? ????? 2101? ?0??? ? Chuandongocoelurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ???0? ????? ????? ????? ????0 ????? ????? ???0? ????? ???0? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 00??? 00111 01011 ???1? 0???? ?0??? ????? ????? 00?10 10001 00000 ?1000 041?2 12101 1???2 1110? 100?? 100?? ? Coelophysis 00000 00000 ??001 10?0? 00000 0?000 10111 01010 0?000 ?0100

bauri 000?0 0000? 00000 00000 01100

00101 ?0000 00?00 01000 10010 02100

21100 01101 00000 00000 00110 00001

0010? 10100 01000 0011? 00011 01111

???0? 000?0 0?000 1100? 01111 00011

01101 00000 00000 01?00 0100? 00000

0100? 0?0?2 01100 00010 10000 0

00000 ??1?? ?1000 02000 00100

00?00 ??0?? 11000 10100 00100

00101 1?0?0 ?0000 0000? 00110

??000 ?0?00 01011 0[01]?00 [01]00?0

Coelophysis rhodesiensis 00??? 21100 0010? ??00? 01101 100?0 00000 00?02 00101 00001 [01]0000 0000? 00010 00000 00?01 10?00 00000 00000 00002 00100 ?000? 10000 00000 10001 00000 00000 00000 00100 00000 01000 00000 01101 1?000 11000 00000 01011 00000 00?00 00000 01000 0?000 0?11? 11000 01?00 00010 02000 10100 0?001 0[01]000 11011 01010 00000 10010 00110 00111 01111 0000? 1000? 01110 00100 00111 [01]0011 02000 ?0100 ?1100 02100 00001 01111 00011 01000 0 Compsognathus 00??? 10000 0100? ???0? 01000 2100? 1??00 00?00 00000 ??000 00?00 0?00? 00??? ?1?00 ????? ???0? ?0??? ?0??0 0???? ?0??0 ?0??? ???1? 1???? 1?000

The phylogeny of Tetanurae 0?00? 0???? 0011? ?1?0?

?0??1 0000? ???0? ?????

00??1 ?1??0 000?? ?0101

0100? ??000 010?1 1????

0000? ?0?00 1??11 ???01

0??00 ?1?0? 0011? ??0?1

00100 0111? 20?20 1?010

295

?100[12] ????? ??100 ?001? ?0011 00?00 01110 0?0?? [12]??0? ??0?? ?0??0 1?131 1?011 ?11?1 0??00 2???1 11??? ????? 1


Concavenator ????? 0???? ???0? ????? 00100 2??0? 1?00? 1111? 10102 01000 0?110 ??10? ?2??? ?1??? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1020? 0???? ??[01]0? ?100? ????? ????? ????? ?[12]?1? ?10?? ??1?? ?0211 ?1?1? ?[01]100 ?000? ???1? 11111 0?10? 1??0? 1??10 ??0?1 01000 1???? ????? ????? 0?0?0 ?10?1 1??11 10110 20??? ????? 02011 ???1? 22??1 100?? ????? ????? ??[12]2? ?0?0? 1?1?? ???0? ??0?1 ?00?? ? Condorraptor ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 00100 01??? ????? 01001 ????? ???00 00?00 101?0 00010 00?20 01?01 11010 01?00 02??? 01??1 01??? ????? ????? ????? ????? ????? ????? ????? ????? 0?1?? 01?01 010?? ????? ?0??? ?0??? 00001 100?? ????? ?0??0 10000 01?00 ??11? ????? ????? ????? 1?01? ??0?? ? Cryolophosaurus ????? ????? ????? ????0 0???? ????? 1??00 ?1001 ?0101 0?000 ?0111 ??0?0 0[01]0?1 ?1000 0?10? ?0000 00??1 00010 00001 ??1?0 010?? ??0?? ????1 ??0?? ????? ????? 10?00 00000 00?0? 01??? ???0? 01000 ????? ???00 ?0?00 0?0?1 000?? 00100 0??01 0?0?? ???00 0200? ???0? 01??? ????? ????? ????? ????0 ?1000 1???? ????? ????? 00??? 00??? 0???? ?011? ?0??? ?0??? 0?0?? 10011 001?0 ?0011 01000 00000 ?2??? ???01 00101 1??0? ????? ????? ? ‘D. sinensis’ ?0?0? 000?0 010?? ????? 00020 2??0? 1??0? ?1013 00100 ??000 ?00?0 0?01? 000?? ?1??0 ????? 1?0?? ????? ????? ????? ????? ????? ????? ????? ????0 0??0? 00??? 1???? ????? 0??0? 0???? ??10? 0100[12] ?1?00 ??000 10000 ?0010 00000 100?0 00001 ??000 ?1000 020?? 01?01 0??01 0001? 00011 21110 00000 01000 1???1 01011 1?000 000?0 101?0 00011 1011? [01]0020 10000 ?2000 10111 00111 00001 01000 ?1000 02200 1010? 0???? 0???1 10011 100?? 1

Dilophosaurus 01?11 21100 00100 0000[01] 00100 01000 00000 ?1002 00100 ??00? 20010 0001? 00010 01000 ?1101 10100 00000 00?00 00002 00101 00000 1??0? ?0??? ??201 10001 0000? 10000 00[01]10 00000 01000 00101 01101 ??000 11100 10000 01011 00000 00000 00101 01000 01000 0201? ?1000 01000 00010 01001 10100 00001 01000 1?111 01100 01000 10010 00110 00011 00110 [01]000? 10000 01000 1?001 00110 [01]0000 00000 00100 02110 02100 00101 01101 00011 000?0 ? Dubreuillosaurus 01??? 100?0 01010 1?000 00100 1[12]000 1??0? ?0??0 ??002 ??001 ?0??? ?1011 ?0010 11?00 0000? ????? ????1 0??20 11100 ?0??1 10010 ??0?? ????0 10?01 10111 ??201 ????? ??000 0000? 01000 00100 ?1001 ????? ????0 ????? ????0 ?00?? 0?1?? ????1 ????? ??1?? 000?1 0???? 0110? ????0 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????1 1???? 0?0?? ????? ????1 ?01?? ????? ????? ????1 ????? ? Duriavenator 00??? 100?? 01010 11000 ?0?0? 120?1 ???0? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????1 10101 ????? ????? ??000 0010? 0??0? 00100 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ? Elaphrosaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????

296


????? 1?000 ????? 01000

M. T. Carrano et al. ????? 00000 ????? 0210?

????? 02111 00011 ?21?0

????? ?0000 10000 ?????

????? 011?? 00?11 ?111?

????? 010?? 11111 001??

????? 0110[12] ????? ???00 10000 0?011 00000 ????? 0???1 1?011 10100 00100 0???? ????? 00?[12]? ????? 0?0?0 00011 00110 ?0011 0??00 ?1??? ?

Eocarcharia ?01?0 02011 ????? ????? ????? ????? ????? ????? ????? ?????

????? 11??? ????? ????? ????? ?????

????0 1???? ??100 ????? ????? ?????

00000 ????? 0010? ????? ????? ?????

11100 ????? 01??? ????? ????? ?????

10100 ????? ??10? ????? ????? ?????

21111 ????? ????? ????? ????? ?

11?0? ????? ????? ????? ?????

????? ????? ????? ????? ?????

????? ????? ????? ????? ?????

????? ????? ????? ????? ?????

????? ????? ????? ????? ?????

Eustreptospondylus ?1??? 20000 011?0 11000 00010 120?1 ???0? ????? ?1002 00001 ????? ?10?1 0001? 1??00 000?? ?0111 10001 0??10 01000 100?1 10000 12??? ????? ??001 10111 ????? ????? ??000 00?0? 01000 00100 11001 ???01 10100 10001 10110 00001 00110 00101 ?100? ?1?00 00??? ?1??? ????? 111?0 ???01 21210 ?00?? ????? ????? ????? ????? ?0011 01011 00011 ?0113 00?2? 1?000 00101 0100? 11111 10000 11000 01000 02111 101?1 10102 111?? 1001? ?000? ? Fukuiraptor ????? ????? 0???? ????? ????? ???11 ????? ????1 1??01 ??12?

????? ????? ????? ????? 0???? ???01

????? ????? ??100 ????? ?10?? 11212

????? ????? 0010? ????? ????? 1??01

????0 ????? 0110? ????? ????? 2?1??

?0?0? ????? ???00 ????? ????? ?0???

????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 110?[12] ????? ???1? ????? ????0 ????0 ????? 12001 21110 1?001 ?1010 ????? ????? ????? ????? 21012 101?1 10001 ?

Giganotosaurus 02??? 000?? 0?00? 11?10 00010 01101 ???10 ?1110 10002 1110? ????? ??202 1211? ?1121 1??0? ?1?00 020?1 10121 ???00 100?0 01121 1?1?? ????1 11??2 01010 ????? 1?111 1110? 00200 00??0 00100 11012 ?01?0 ??111 001?0 1?1?0 0?100 ?1100 1?21? 2???? ?2?00 000?? ?1??0 ?10?? 0???0 1?0?? ????? ????? ????? ????? ????? ????? 00?11 01001 02111 10110 20120 1?120 02011 11000 22011 101?? 1?0?? ?1001 ?3122 1010? 10??? ????? ????? ????? ? Irritator ????? 2???? 02?0? ????? ?0110 0200? 10001 0??00 00010 ?2011 10010 000?? 00011 ?1100 0???? 0?0?? ?11?1 0??10 ?1001 00010 21000 ?101? 10??? ??0?? ????? ????? 10?11 ?0??? 1101? 19??? ???1? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1???? ????? ????? ????? ????? ????? ????? ????? ????? ? Leshansaurus ????? ????? ?1?1? 1?001 ??1?0 [01]???1 ????? ????? ????? ????? ????? ????? ????? ?1000 0???? ????? ????0 00000 01002 1001? ?00?? ??0?? ????? ????? ????? ????? ????? ???00 ???0? ????? ??[01]0? 1100? 1?001 0??01 10000 121?0 000?? 0???? 00000 1100? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1???? ????? ???0? 0???? ??011 0?011 1011? [12]0??? ????? 0??0? ????? [01]???? 10000 1???1 ?10?? ?2111 10?0? ????? ????? ????? ??0?? ? Lourinhanosaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1100? ????? ???0? ????? 1?1?0 00?00 0?0?0 0???1 ?1?0? ?1100 000?? ???0? 111?? ????? ????? ????? ????? ????? ????? ????? ????? 00?01 010?1 01111 1011? 2???? ?1??? 0?001 100?? 11011 101?0 110?? ???01 ?21?2 101?? ????? ????? ????? ????? ? Magnosaurus ????? ????? ????? ????? ????? ????? ???0? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????1

The phylogeny of Tetanurae 10?11 ????0 ????? 01000

????? ????? ????? ?21??

????? ????? ?0??? ???0?

??000 ???00 01??1 11???

0000? ????? ????? ?????

0???? ????? ????? ?????

????0 ????? ????? ?????

297

???0? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ??0?? 0???? ????? 1???? ???0? 110?0 ?


Majungasaurus 02101 00010 00000 10000 00010 01000 00111 20?10 02000 0?101 20000 00202 02012 ?1111 01101 10000 10001 0100? 00000 010?0 1101? 10000 00010 10110 01010 01100 10001 00101 00000 00100 00100 01102 10011 00100 11000 12110 00000 10100 00001 021?1 11100 1201? 01000 011?1 011?1 0?011 10200 0010? ????? 0???? ????? ???0? ?0011 10000 00011 11111 ????? ?0??? 1???? ????? 0???? 10011 0?100 01010 02102 11111 11102 1111? 0001? ?1101 ? Mapusaurus ????? ?0??? 00000 11110 00010 01101 ?1010 11110 10002 1110? ?1111 102?2 12112 1???? ????[02] 01?00 0?0?? ????? ????? ????? ????? ????? ????? ??002 01010 ?0100 1???? ??100 00200 00?0? ??100 11012 ???01 ???1? ?0?00 10??0 001?? 01?00 10201 2???? ?2100 00??? ????? ?1?11 012?0 ????1 ???10 ?10?? ????? 0???? 1???? ????0 00??? 0?0?1 02?11 10110 [12]???? ????? 0?011 ?1000 22??1 10111 1?00? 1100? ?3122 10101 10??2 11101 2101? ?0010 ? Marshosaurus 000?? 00000 00010 01000 11020 11001 ?0000 10?00 0???? ????? ?0?10 01?10 000?? ??100 0000? 10?0? 01?01 00020 00000 1001? 0?00? 1?0?? ????? ????1 0001? ?0??? 1?011 ??001 00100 01000 00100 11001 ?0111 11000 10000 12110 00000 00120 01101 ?1??? ????? ????? ?10?? ????? 0?1?? ????? ????? 1???? ????? ????? ????? ????? 00?11 01001 01011 ?0110 01020 1?000 0?001 11000 ????? 1???? ????? ????? ????2 1???? ???0? ????? ????? ????? ? Masiakasaurus 0???? ?0??? 00100 10000 01000 01000 ???0? ????? ???10 ????? ????? ?0??2 0101? 10??? ????? ?0?00 1??00 ????0 00?00 01??0 0???0 100?? ????? ??111 01000 01190 ??00? 00010 00000 00??? 00200 01102 ?0011 00100 11000 111?0 00000 10100 00000 02011 11000 1011? 01000 01?01 01111 10011 10100 001?? ????? ????? ????? ???00 1001? 10000 00011 ?1111 11120 10000 10010 01??1 00110 10011 02100 01010 02102 ?1111 11202 1111? 0011? ?1101 ? Megalosaurus ????? ????0 000?0 11011 00?00 110?1 ?000? ????? ????? ????? ?1?11 1???? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????0 00101 ????? 0?011 00001 0010? 01??0 0?100 ???01 ????? ???0? ????? 1???? ?0?1? ???10 00201 11000 01?00 00??? 010?? ???01 110?0 00001 21211 0?000 01000 0???? ????? ????? 00101 01011 01011 10113 ?00[12]? ????? 0?10? ?1?00 11101 10000 1?000 01000 0221? ?0?0? 1???? 1???? 1001? ?00?? ? Megaraptor ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 11012 ????? ???11 ?0100 121?0 ????? ???1? ????? ????? ???11 0?0?? ????? 1?111 011?0 121?? ????? ????? 01010 11?2? 01011 11111 0???? ????? ????? ????? ???2? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 2?11? ????? ? Metriacanthosaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?1001 ????? ????0 ??0?? 1?1?0 0011? 00?00 01211 1???? ?1?00 0?0?? ?1??? ?1??? ????? ????? ????? ????? ????? ????? ????? ????? 00??1 01001 01111 ?011? ?012? 1?0?0 0?101 ?01?1 11??1 10?11 11000 1??0? ??21? ????? ????? ????? ????? ????? ? Monolophosaurus 001?1 000?0 0100? ????? 00000 2000? 10001 11013 00100 ?9001 2[01]110 00110 010?1 11100 01?02 10000 00??1 00010 00000 10??? ??0?? ??01? 1????

298 ??001 00000 ????? ?????

M. T. Carrano et al. 00001 0?100 ????? ?????

00201 00101 ????? ?????

00?0? ?10?0 00110 ?????

??00? ?1?00 00101 ?????

0010? 0?00? 01011 ?????

0??00 010?? 1011? ?????

00101 11001 0?001 00100 10?00 10110 ????? ????? ????? ????? ????? ????? 00?[12]0 100?? ?0001 000?0 ????? ????? ????? ?


Neovenator 00??? 10000 01000 10101 00000 21001 ???10 11011 0???? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????1 ????? ????1 10000 ????? ????? ??100 00100 01?02 00100 11012 ?1??? ??111 10100 12110 00100 01101 01011 0?0?? ?2100 001?? ?1?10 ?10?1 011?0 120?? ????? ????? ????? ????? ????? ????? 01?11 01001 02211 10110 20120 1?120 00001 ?1011 020?1 10111 10001 11001 13122 1010? 10??? 1???? 21011 1?010 ? Ornitholestes 000?? 000?0 0000? ????? 00000 2100? 1?000 01000 00101 0?001 2?110 00001 000?1 ??100 01?02 0?00? ?0??1 000?0 0???0 ????0 ??0?? ????? ????? ??000 00?0? ?0?0? 0?010 0111? 00000 0??00 00100 1100[12] ????? ????0 10010 1?0?0 00?00 01000 00001 0100? ?1?00 010?? 010?? 11??? ????? ??001 11100 00000 0???0 1??11 ??111 ??1?? 00?00 01011 12111 10110 20??? ????? 01001 00000 21??2 11??0 10000 1???? ???2? ?0?0? ????? ????? 2001? ?0??? ? Piatnitzkysaurus ????? ????? 000?0 10000 10020 110?1 ???0? ????? ????? ????? ????? ????? ????? ?1??? 0???? ????? ????1 0?010 00000 10?11 01000 ?10?? ????? ????2 010?1 ????? ????? ??001 0010? 01??? ??000 01001 ?0011 11000 10000 12110 00010 00120 0?101 11000 01?00 0?0?? 010?? ???01 011?1 12001 21110 1100? ?100? 1???? ????? ????? 001?1 01001 01011 ?011? 0002? 10000 00001 10000 11?11 10000 10000 01000 02212 1010? 10??? 1???? 1001? ?0??? ? Piveteausaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????0 11?00 0???? ????? ????1 000?0 ?1002 100?? 1002? 110?? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ? Poekilopleuron ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ??100 ?0??1 ????? 111?? ????? ???01 2??10 11000 10000 01??? ?0?11 ????0 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?10?? ?2??? ?0?01 11102 1?10? ????? ??000 ? Proceratosaurus 000?? 000?0 0100? ????? 00000 2100? 1??0? ????? ??0?? ????? 20110 0000? ????? ????? ?1??2 0??0? ?0??? ????? ????? ????? ??0?? ????? ????? ??00[02] 00?0? ?0??? 00??1 ?1?11 0000? 0??00 00000 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ? Saurophaganax ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ??0?? ????? ????? ????? ????? 0???? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1100? 1???? ????? ?0?00 1?1?0 00?0? ?0??? ???11 ????? ???00 ??0?? ????? ?10?? ????? ???01 2?010 1100? ????? ????? ????? ???00 001?1 01001 0???? ????? 20?[12]0 1?1?0 0??01 ????? 210?1 10??0 1???0 11001 ?21?? ????? ????? ????? ?00?? ????? ? Shaochilong ????? ????? 000?? 11?10 10010 01101 11?00 ???1? ????? ????? ????? ????? ????? ?1?01 1???? ?0?00 02001 1???1 ?0000 100?0 011?? 120?? ????? ?????

The phylogeny of Tetanurae ????? ????? ????? ?????

????? ????? ????? ?????

????? ????? ????? ?????

??1?? ????? ????? ?????

????? ????? ????? ?????

????? ????? ????? ?????

????? ????? ????? ?????

299

????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?


Shidaisaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ???0? ?1100 0???? ????? ????1 10?0? ????? ????0 ????? ????? ????? ????? ????? ????? ????? ????? 0??0? 00??? ????? ???0? ?0110 000?? ????? 1???? 001?0 0???? 01211 01000 01?00 ????? ??0?? ????? ????? ????? ????? ????? ????? ????? ????? ????? 00??0 ?1??1 ???11 10110 20?[12]0 ??00? ?20?? 0??0? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ? Siamotyrannus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ???0[12] ????? ????? ????? ????? ????? ????0 ???11 01000 ?0?00 0???? 010?? ????? ????? ????? ????? ????? ????? ????? ????? ????? 00201 01001 02?11 10112 20121 10000 ??101 101?? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ? Sinraptor dongi 00001 00000 00000 00000 01000 21111 11000 11010 00002 11000 01110 00100 01001 ?1100 00112 00100 02001 10110 00100 10010 01120 10011 1?111 11000 00011 00200 11111 00000 0010? 0?100 00100 11001 10110 00100 10000 12110 00110 00110 01211 010?0 ?1?00 0?000 010?? ?1?01 01110 0???? ????? ????? ????? ???2? ????? ?11?0 00201 01001 01011 10112 20121 11000 02101 10101 11011 10111 10000 11000 02212 10101 10102 11101 21011 10010 1 Sinraptor hepingensis 00001 00000 0000? ????? 01000 2111? 1?000 11011 00002 11000 01110 00100 01001 ?1100 00112 00100 0???1 1?120 00?00 ????? 01??? ???1? 1???? ??000 0001? 00?00 1???1 ?000? 00?00 0?100 00100 11001 10110 00100 10000 12110 00110 001?0 01211 ?1100 01100 0100? 0101? 11001 01110 0?0?? ????? ????? ????? ????? ????? ????? 00001 01001 01?11 10112 [12]0021 11010 02101 10?10 110?1 10111 1?00? ????? ????? ????? ????? ????? ????? ????? ? Spinosaurus ????? ????? 10?0? ?011? 0???0 ?0201 ????? ????? ????? ?????

110?1 ????? ????? 0?00? ????? ?????

20101 ????? ??010 ???00 ????? ?????

1210? ????? 11011 0?0?? ????? ?????

???0? ????? 18013 0???? ????? ?????

?0000 ????? 11?10 ????? ????? ?????

020?? 1??0? ????0 ????? ????? ????? ????? ????? ????? ????? ????? ??001 1100[12] ????? ????? 1000? ?21?0 100?0 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?

Streptospondylus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1100[12] ????? ???0? ?0?01 ??1?0 00001 0?1?0 ???01 0???? ???00 0?0?? 0?0?? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ??0?? 1?000 ????? ???0? ????? ????0 1000? 01??? ?2??? ????1 10102 111?? ????? ????? ? Suchomimus 11110 20101 12101 ?0?00 00010 0200? 1??0? ????0 ????? ????? ????? ????? ????? ?1??? ????? ???11 0110? ????? ????? ????? ????? ????? ????? ????1 10??? ???1? ????? ??010 01011 0?013 00101 1100[12] ????? ??100 ?0001 121?1 11010 00110 00201 ?10?0 ???00 000?? ?1001 ?0001 011?0 10001 21111 00010 01101 0???? ????? ???10 0011? 010?1 00011 10110 00?[12]? 0?00? 00001 1?00? 11??1 10100 11010 0100? ?4121 1010? 10202 1??0? 1???? ????? ? ‘S. zigongensis’ ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????

300 ????? 0?1?0 00111 0?00?

M. T. Carrano et al. ????? ????? ????? 00?0? ?1??? ??00? 1100[12] ????? 0010? ?0000 12110 00100 0?011 ?1??0 ??000 ?100? ????? ???01 011?0 ???01 10210 01000 01000 1??2? 111?? 00001 010?1 0??11 10110 101[12]0 ?100? 02001 10111 11?1? ?0??1 10?00 ?211[12] 10?0? ?010? ????? ????? ????? ?


Torvosaurus 00??? 100?0 010?? 11011 00010 12000 ??00? ???00 ?1012 00001 ?0111 11011 000?[01] ????? ????? 00111 1?01? ????? ????? ????? ????? ????? ????? ????0 10?0? ????? ????? ??000 00100 01001 00100 11011 01021 00100 10001 12110 00011 00110 00101 11000 01110 000?? 010?1 01001 11000 00001 21211 00000 0100? 0[12]??? 00110 11010 00001 01011 01011 10113 000[12]? 10000 02001 01000 1110? 10000 11000 01000 02121 10101 10102 1110? 1001? ?0??? ? Tyrannotitan ????? ????? ????? ????? ????? ????? ???1? ????? ????? ????? ?1111 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????2 0?010 ?0??? ????? ??1?? 0?20? 0???? ????0 1101? 1???? ???1? ??100 1?1?0 0??00 ?1?00 1?2?1 ????? ??000 ??0?? ????? ???11 ????0 1???? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?0??? ?0?20 1???? ?20?? ???0? 220?1 ?0111 1?0?0 1???? ???2? ????? ????? ????? ????? ??010 ? Xuanhanosaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1100? ????? ???0? ????? 1???? ??000 ?0??0 0??11 ????? ??1?? ????? ????? ???0? 1???1 ?200? 2111? 01000 01000 1102? 00110 1?100 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ? Yangchuanosaurus 0000? 00000 0000? ????? 00000 2111? 1?000 11011 00002 ?1000 01110 0010? 010?1 ?1??0 ?0?12 0???0 ?0??? ????? ????? ????? ????? ??011 1???1 ??000 00?1? 00??? 1???1 ?0??? 00?0? 0??00 00100 1100[12] ?0110 001?0 10000 12110 00100 0?1?0 00111 01000 0?000 0100? 010?? ????? ????? ????? ????? ????? ????? ????? ????? ????? 00000 01001 01?11 10110 10120 ?100? 02001 1?111 11??0 101?1 10??? ?100? ?211? ?0101 10102 1??0? ????? ????? ? CV00214 ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? 1100[12] ?0110 001?? 10000 12110 00100 0???0 00[12]11 ??0?? ???00 ?1??? ????? ????? ????? ???01 2010? 0?001 01000 ??0?? 000?? ????? 000?0 010?1 0???? ?0110 [12]0?[12]0 ??00? 0?001 ???1? [12]1?10 111?1 10?00 ???0? ??1?? ?0?0? 1???? ???01 ????? ??0?0 ?