the phylogeny of tetanurae (dinosauria: theropoda)
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The phylogeny of Tetanurae (Dinosauria: Theropoda)Matthew T. Carrano a , Roger B. J. Benson b & Scott D. Sampson ca Department of Paleobiology, National Museum of Natural History, Smithsonian Institution,Washington, DC, 20560, USAb Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge, CB23EQ, UKc Natural History Museum of Utah and Department of Geology & Geophysics, University ofUtah, 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
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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
aDepartment of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA; bDepartmentof 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
(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 extantAves. 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) tetanuransled to uncertainties regarding their basic interrelationships. This in turn prevented determining the relationships of manyincompletely known taxa that nonetheless document a global radiation spanning more than 120 million years. We undertookan exhaustive examination of all basal tetanurans and all existing character data, taking advantage of recent discoveriesand adding new morphological, temporal and geographic data. Our cladistic analysis of 61 taxa achieved significantlyimproved 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 basalmostclade (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 formsand hundreds of additional reported tetanuran occurrences. Tetanuran evolution was characterized by repeated acquisitionsof giant body size and at least two general skull forms, but few variations in locomotor morphology. Despite paralleldiversification of multiple lineages, there is evidence for a succession of ‘dominant’ clades. Tetanurae first appeared by theEarly Jurassic and was globally distributed by the Middle Jurassic. Several major clades appeared prior to the breakup ofPangaea; as such their absence in specific regions, and at later times, must be due to poor sampling, dispersal failure and/orregional extinction. Finally, we outline a general perspective on Mesozoic terrestrial biogeography that should apply to mostclades that appeared before the Late Jurassic.
Keywords: evolution; systematics; Mesozoic; taxonomy; morphology, biogeography
Introduction
Theropod biology, evolution and phylogeny have receiveddisproportionate attention within the study of dinosaurpalaeontology. Currently, non-avian theropod taxacomprise 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 ofan unusually diverse dinosaur clade, but this perceiveddiversity is heavily mitigated by certain anthropogenicbiases. Certainly the inherently dramatic predatory qual-ities of these organisms have lent them a special focus,but significant research efforts have also been directedat resolving the problems surrounding bird origins (e.g.Gauthier 1986; Turner et al. 2007). As a result, theropodsare probably both oversampled and overstudied relative toother dinosaur groups.
In spite of this, many individual theropod taxa are aspoorly studied as most other dinosaurs. Because research
∗Corresponding author. Email: [email protected]
has tended to focus on the especially impressive (e.g.Tyrannosaurus) and the nearly avian (e.g. Dromaeosauri-dae) amongst the Theropoda, much less has been learntabout the 100+ more primitive forms. Indeed, those aspectsmost fundamental to understanding the evolution of basaltheropods – their phylogenetic interrelationships – haveonly recently received much specific attention (e.g. Serenoet al. 1996, 1998; Holtz 2000, 2004; Rauhut 2003; Carrano& Sampson 2008; Benson 2010a).
Most previous studies of basal tetanurans have focusedon placing individual, usually new, taxa into a phylogeneticcontext (e.g. Sereno et al. 1994, 1996) and tend to includemore complete forms in the hope of achieving the great-est resolution. By contrast, other works have attempted toresolve relationships amongst theropods as a whole (e.g.Holtz 2000; Rauhut 2003). In general (see below) thesehave produced more limited resolution, often conflicting insignificant details with one another. This probably arisesfrom a paucity of characters constructed specifically to
ISSN 1477-2019 print / 1478-0941 onlineCopyright C© 2012 The Natural History Museumhttp://dx.doi.org/10.1080/14772019.2011.630927http://www.tandfonline.com
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resolve basal tetanuran relationships (rather than particu-lar ingroups of closely related taxa), which itself resultsfrom the historical lack of focused study.
Several years ago, two of us (MTC and SDS) undertooka major project to address these difficulties. We began witha comprehensive review of theropod interrelationships,focusing on all taxa more primitive than Coelurosauria. Thiswork has involved first-hand examination of hundreds oftheropod specimens, critical review of nearly 700 existingcharacters from all published studies, and inclusion of manytaxa and characters never previously incorporated into anyphylogenetic analyses. RBJB independently undertook areview focused on non-coelurosaurian (‘basal’) tetanuransand also reviewed existing taxa and character descriptions.The present work represents a combination of our efforts.
This paper is the sixth documenting our phylogeneticresults. The first (Carrano et al. 2002) provided evidencethat Ceratosauria was a lineage independent of Coelo-physoidea and more closely allied with Tetanurae. Thesecond (Carrano et al. 2005) produced a phylogeny ofCoelophysoidea along with a redescription of the smalltheropod Segisaurus. The third (Carrano & Sampson 2008)focused on the ingroup relationships of Ceratosauria.Benson (2010a) focused on resolving the relationships ofMiddle Jurassic theropods, including the problematic Euro-pean ‘megalosaurs’, and we recently recognized a distinctlineage of Cretaceous allosauroids (Benson et al. 2010).
These works complement several descriptive studies ofindividual theropod taxa, including the ceratosaurs Majun-gasaurus (Carrano 2007; Sampson & Witmer 2007) andMasiakasaurus (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 mega-losaurs Duriavenator, Magnosaurus and Megalosaurus(Benson 2008a, 2010a, b), the carcharodontosauriansChilantaisaurus, Kelmayisaurus, Neovenator, and Shaochi-long (Benson & Xu 2008; Brusatte et al. 2008, 2010b,2011), and the coelurosaur Stokesosaurus (Benson 2008b).Together these have provided new anatomical and characterdata for the present study.
Here we present a systematic analysis with the goal ofproducing a well resolved, highly documented and compre-hensive phylogeny of all basal (= non-coelurosaurian)tetanurans. Several non-tetanurans and coelurosaurs areincluded as reference taxa to define the basic interre-lationships of the major theropod clades; ceratosaursare thus included but at a reduced level of sampling.Although we employ species level sampling and avoidusing suprageneric groups, we have excluded a number offragmentary forms in an attempt to achieve some measureof resolution. When possible, we have used criteria forSafe Taxonomic Reduction, omitting only those highlyincomplete taxa whose character combinations are notunique (Wilkinson 1995). Additionally we employ Adamsconsensus methods (Adams 1972) to identify ‘wildcard’
taxa a posteriori. This is a manual implementation of thestrict reduced consensus method (e.g. Wilkinson 2003).
Historical background
Constituency and placement of TetanuraeAlthough the term is commonly used today in dinosaurstudies, Tetanurae was only recently recognized and named(Gauthier 1986). Gauthier’s study is an important bench-mark for theropod systematics as it represents the earliestcomprehensive cladistic treatment of Theropoda and thesource of much of the basic topology on which currentstudies are still constructed.
That said, the taxa now called tetanurans include theearliest named dinosaur (Megalosaurus Buckland, 1824),and a wide array of forms discovered since that span nearlythe entire temporal and geographic range of Dinosauria.As such, these taxa have been of interest to theropodand dinosaur workers since the 19th century, even thoughthey have rarely been recognized as belonging to a singleevolutionary lineage. Indeed, within the ranks of Tetanu-rae are found some of the most problematic taxa in allDinosauria.
Megalosaurus has achieved an almost mythic statusamong these. A fragmentarily known form from the MiddleJurassic of England, all known topotypic specimens aredissociated and were deposited allochthonously in marginalmarine sediments. Its early discovery and description(Buckland 1824) predate both the term Dinosauria andany standardization of palaeontological reporting, therebycontributing to the confusion surrounding its identity andrelationships. Unfortunately, as a hallmark theropod, Mega-losaurus served as a taxonomic ‘attractor’ for decades,resulting in the unstable, often incorrect, and usually unjus-tified placement of dozens of referred species within thegenus. This is discussed in further detail below.
For a century after the description of Megalosaurus,most large carnivorous dinosaurs were grouped into thefamily Megalosauridae within the order Theropoda (e.g.Baur 1891; Hay 1902; Romer 1956; Walker 1964; Fig. 1).At times distinct European and North American lineageswere recognized, with the latter often referred to as Dryp-tosauridae (Marsh 1895). Occasionally, workers separatedindividual taxa into discrete families if their morphologyappeared aberrant (e.g. Labrosaurus, Labrosauridae;Ceratosaurus, Ceratosauridae; Spinosaurus, Spinosauri-dae), but Megalosauridae remained the only multi-taxicgroup of large theropods employed consistently duringthis time. In addition, nearly all researchers arrayedthe families of carnivorous dinosaurs serially withinTheropoda (or Saurischia), with only a few forms segre-gated into taxa of higher rank (e.g. Ceratosauria, Coeluria,Compsognatha).
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The phylogeny of Tetanurae 213
Huxley 1870
Dinosauria Iguanodontidae Megalosauridae Euskelesaurus Laelaps Megalosaurus Palaeosaurus Poekilopleuron Teratosaurus Scelidosauridae
Lydekker 1888
Theropoda Anchisauridae Coeluridae Compsognathidae Megalosauridae Aristosuchus Bothriospondylus Dryptosaurus Megalosaurus Zanclodon
Marsh 1895, 1896
Theropoda Ceratosauria Compsognatha Hallopoda Coeluria Anchisauridae Dryptosauridae Allosaurus Coelosaurus Creosaurus Dryptosaurus Labrosauridae Labrosaurus Megalosauridae Megalosaurus Plateosauridae
Zittel 1911
Theropoda Anchisauridae Coeluridae Compsognathidae Labrosauridae Labrosaurus Megalosauridae Allosaurus Antrodemus Coelosaurus Creosaurus Dryptosaurus Laelaps Megalosaurus
Hay 1902
Theropoda Anchisauridae Ceratosauridae Coeluridae Hallopidae Megalosauridae Allosaurus Antrodemus Coelosaurus Creosaurus Deinodon Dryptosaurus Megalosaurus Palaeoctonus Suchoprion Troodon Zapsalis Zatomus Ornithomimidae
Huene 1909
Saurischia Ceratosaurus Coeluridae Compsognathidae Megalosauridae Allosaurus Antrodemus Creosaurus Labrosaurus Megalosaurus Plateosauridae Sauropoda Sellosauridae Thecodontosauridae Zanclodontidae
Huene 1914dCoelurosauriaPachypodosauria Ammosauridae Massospondylidae Megalosauridae Plateosauridae Sauropoda Sellosauridae Zanclodontidae
Huene 1923b
CoelurosauriaCarnosauria Megalosauridae Altispinax Antrodemus Megalosaurus Streptospondylus Cladeiodon Dryptosaurus Erectopus Spinosaurus Teratosaurus Zanclodon
Williston 1878
Dinosauria Ornithoscelida Allosaurus Compsognathus Creosaurus Dryptosaurus Hypsilophodon Iguanodon Laosaurus Nanosaurus Poekilopleuron Sauropoda
Marsh 1882
Theropoda Amphisauridae Compsognatha Dryptosauridae Dryptosaurus Labrosauridae Labrosaurus Megalosauridae Allosaurus Coelosaurus Creosaurus Megalosaurus Zanclodontidae
Gilmore 1920
Theropoda Ceratosauridae Coeluridae Megalosauridae Antrodemus Deinodon Dryptosaurus Labrosaurus Tyrannosaurus Ornithomimidae
Baur 1891
Megalosauria Anchisauridae Coeluridae Compsognathidae Dryptosauridae Dryptosaurus Megalosauridae Ceratosaurus Megalosaurus Zanclodontidae
Huene 1926bHuene 1926a
CoelurosauriaPachypodosauria Carnosauria Megalosauridae Altispinax Erectopus Megalosaurus Prosauropoda Sauropoda
CoelurosauriaCarnosauria Megalosauridae Antrodemus Megalosaurus Spinosauridae Spinosaurus
Huene 1959
CoelurosauriaCarnosauria Allosauridae Dinodontidae Teratosauridae
Huene 1929
CoelurosauriaPachypodosauria
Carnosauria Dinodontidae MegalosauridaeProsauropodaSauropoda
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
Plateosauridae Euskelosaurus Gresslyosaurus Pachysaurus Plateosaurus Poikilopleuron Sellosaurus Teratosaurus Zanclodon
Figure 1. Historical classification schemes that have included theropods which are now classified as tetanurans, 1870–1956. Taxa recoveredas non-coelurosaurian tetanurans in the present study are indicated in bold.
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In the early 20th century, Huene produced several reviewsof the large theropods then known, resulting in a seriesof increasingly refined classifications. He was primarilyconcerned with the overall systematics of Saurischia, atopic he worked on for nearly 50 years (e.g. Huene 1914a,b, c, d, 1920, 1923, 1926a, b, 1932, 1956). He describedmany new taxa, some of which are still considered valid,but his most significant achievement was the creation ofan underlying systematic scheme for predatory dinosaursthat effectively replaced the serial arrangement of previousdecades.
Initially Huene (1914a, b, c, d) arranged the small,lightly built forms into the infraorder Coelurosauria andput the larger bodied taxa into the infraorder Pachypo-dosauria, although still within the family Megalosauridae(Pachypodosauria included many forms now consideredto be prosauropods and/or basal sauropodomorphs). Later,however, Huene (1920) transferred the large, strictlycarnivorous taxa from the Pachypodosauria to a newinfraorder, Carnosauria. The remaining pachypodosaurswere allied with sauropods in the Sauropodomorpha.Carnosauria (Huene 1923, 1926a, b) then included allknown large bodied predators other than Ceratosaurusand tyrannosaurids (Albertosaurus, Gorgosaurus, Tyran-nosaurus). The former was retained within Coelurosauriaon the line leading from Coelophysis; the latter were consid-ered to be closely related to ornithomimids. Althoughboth were placed in Carnosauria by many subsequentworkers, the arrangement posited by Huene (1926a, b) isremarkably close to our current perception of theropodrelationships.
A few workers presented somewhat different viewsduring this time. Unlike Huene, Matthew (1915) retainedlarge theropods in Theropoda, apart from Coelurosauria.Likewise, Nopcsa (1923) separated the large theropodsfrom pachypodosaurs into a separate Megalosauria.Although he retained a distinction between them andsmaller theropods (Coelurosauria), the three groups wereof equivalent rank within Saurischia, alongside Sauropoda.Neither of these had the lasting influence of Huene’sschemes, nor did any of the other variations producedby different workers. But nearly every study empha-sized the phylogenetic differences between large andsmall theropods, and placed basal tetanurans among theformer.
The size-based distinction between carnosaurs andcoelurosaurs rooted in Huene’s classification had a longlasting effect on theropod systematics; for more than halfa century, few workers presented serious challenges tothis organizational scheme. Even the later compilations ofRomer (1956, 1966) and Steel (1970) retained this basicstructure (Fig. 2). Aside from acceptance of Spinosauri-dae and Allosauridae as separate families (e.g. Colbert &Russell 1969; Carroll 1988), little changed for nearly 60years. Among traditional studies, only Kurzanov (1989)
erected new families of large theropods (Streptospondyl-idae and Torvosauridae) within the Carnosauria. Walker’s(1964) study separated megalosaurids and spinosaurids intodifferent superfamilies, and thereby provided more struc-ture than previous classifications, but still maintained bothwithin the Carnosauria. Not until Gauthier (1986) funda-mentally reorganized theropod interrelationships were anylarger changes established. And although Gauthier retainedthe terms Carnosauria and Coelurosauria, as well as theduality of their relationship, his redefinitions effectivelyreplaced earlier size based concepts.
Modern phylogenetic analyses generally agree on amonophyletic Tetanurae that includes a series of generallylarge bodied basal forms that lie outside a monophyleticCoelurosauria (Figs 3–5). Tyrannosauridae is now univer-sally included within Coelurosauria (Novas 1991a; Holtz1994a), whereas ceratosaurs and coelophysoids are basalto Tetanurae. Although some workers prefer the termCarnosauria 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 bodiedtheropods. However, its use as the name for a possibleclade comprising Allosauroidea and Megalosauroidea(Spinosauroidea) as employed by Rauhut (2003) isfaithful to Huene’s (1920, 1923a, b, 1926a, b) originaldefinition.
Ingroup relationships of TetanuraeThe interrelationships of individual tetanuran taxa havebeen even more problematic than their basic organization.The prevalence of serially arranged families within mostearly works (e.g. Marsh 1882, 1895; Hay 1902) often meantthat within-family relationships were paid less attention.
Alternatively, Huene (1923a, b, 1926a, b) and otherstended to describe specific evolutionary lineages oftheropods, in which particular taxa were ancestral to others.There is some analogy to the sister-taxon concept of moderncladistic discussion only in the sense that particular lower-level affinities are implied between individual taxa.
Initial cladistic studies of tetanuran theropods (Paul1984; Holtz 1994a, 2000; Charig & Milner 1997)generally supported a primitive grade of ‘megalosaurs’(Eustreptospondylus, Megalosaurus, Torvosaurus) thatwere arranged as serial outgroups to a small clade ofallosaurs (minimally Allosaurus and Acrocanthosaurus),followed by the Coelurosauria. The consensus was that‘megalosaurs’ represented basal tetanurans, and thus taxamore derived than ‘ceratosaurs,’ but lacked synapomor-phies that might support their monophyly (Fig. 3).
Subsequently, most workers have recognized that manyof these basal tetanurans formed a true clade, usually termedSpinosauroidea (originally Torvosauroidea [Sereno et al.1994; Olshevsky, 1995] but correctly Megalosauroidea[Holtz et al. 2004; Benson 2010a]). Current disagreementscentre on whether this clade is basal to the Allosauroidea
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The phylogeny of Tetanurae 215
Walker 1964
Theropoda Coelurosauria Carnosauria Megalosauroidea Megalosauridae Antrodemus Bahariasaurus Carcharodontosaurus Dryptosaurus Eustreptospondylus Indosaurus Megalosaurus Metriacanthosaurus Tyrannosauroidea Ornithosuchidae Spinosauridae Acrocanthosaurus Altispinax Spinosaurus Tyrannosauridae
Colbert 1964
Theropoda Coelurosauria Carnosauria Megalosauridae Tyrannosauridae
Romer 1966
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
Colbert & Russell 1969
Theropoda Coelurosauria Carnosauria Allosauridae Acrocanthosaurus Allosaurus Megalosauridae Ceratosaurus Eustreptospondylus Spinosauridae Tyrannosauridae
Theropoda Coelurosauria Carnosauria Dinodontidae Megalosauridae Acrocanthosaurus Altispinax Antrodemus Ceratosaurus Megalosaurus Metriacanthosaurus Spinosaurus
Swinton 1970 Barsbold 1983
Theropoda Coelurosauria Carnosauria Allosauridae Ceratosauridae Megalosauridae Tyrannosauridae Spinosauridae
Russell 1984
Theropoda Coelurosauria Carnosauria Allosauridae Allosaurus Aublysodontidae Ceratosauridae Dryptosauridae Megalosauridae Acrocanthosaurus Marshosaurus Stokesosaurus Torvosaurus Tyrannosauridae
Welles 1984
Theropoda Ceratosauridae Coelophysidae Coeluridae Compsognathidae Dromaeosauridae Halticosauridae Marshosaurus Megalosauridae Allosaurus Coelosaurus Eustreptospondylus Megalosaurus Teratosaurus Procompsognathidae
Kurzanov 1989
Carnosauria Abelisauridae Allosauridae Acrocanthosaurus Allosaurus Chilantaisaurus Compsosuchus Marshosaurus Ornithomimoides Piatnitzkysaurus Szechuanosaurus Megalosauridae Gasosaurus Iliosuchus Magnosaurus Megalosaurus Metriacanthosaurus Piveteausaurus Sarcosaurus Yangchuanosaurus
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
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 inbold.
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Megalos
aurus /
Torvo
sauru
s /
Poe
kilople
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Paul 1984
Procom
psogn
athus
Allosa
urus
Eustre
ptosp
ondylu
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Yangc
huanos
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Ceratos
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Coelop
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Ornith
olesti
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tyran
nosau
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maeos
aurs
Piatnitzkys
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Eustre
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PROTOAV
IA
Tyra
nnosau
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Coelop
hysid
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Abeli
saurid
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Cerato
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COMPSOGNATHA
Megalos
aurus
Acroca
nthosauru
s
Allosa
urus
Baryony
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Spinos
aurus
Metriaca
nthosauru
s
Marshos
aurus
Gasos
auru
s
Chilantaisa
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Paul 1988a
Novas 1992
COELUROSAU
RIA
CERATOSAU
RIA
Piatnitzkys
aurus
CARNOSAURIA
Comps
ogna
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Eustre
ptosp
ondylu
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Eorapto
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COELUROSAU
RIA
Spinosaurid
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Torvo
saurid
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Sinrapto
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Allosa
uridae
Afrove
nator
CERATOSAU
RIA
Herrera
saurus
Sereno et al., 1994
Russell & Dong 1993
Allosa
urus
Baryony
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Yangc
huanos
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Trood
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ae
Dromae
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ridae
Ornith
omim
idae
Ovirap
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Coelop
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COELUROSAU
RIA
CERATOSAU
RIA
Torvo
sauru
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Acroca
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Allosa
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Holtz 1994a
Megalos
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Coria & Salgado 1995
Giganoto
sauru
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COELUROSAU
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CERATOSAU
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Carcharod
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Allosa
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Monolo
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Cryolop
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s
Sereno et al., 1996
Figure 3. Results of previous phylogenetic analyses of Tetanurae, 1984–1996. Taxa recovered as non-coelurosaurian tetanurans in thepresent study are indicated in bold.
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The phylogeny of Tetanurae 217
Sereno et al., 1998
NEOTETA
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CERATOSAU
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Spinos
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Irrita
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Suchom
imus
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Sinraptor
P.? va
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COELUROSAU
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Allain 2002
Charig & Milner 1997
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Figure 4. Results of previous phylogenetic analyses of Tetanurae, 1997–2004. Taxa recovered as non-coelurosaurian tetanurans in thepresent study are indicated in bold.
+ Coelurosauria clade (usually called Avetheropoda orNeotetanurae; Paul 1988a; Sereno et al. 1994, 1996,1998; Holtz 2000; Holtz et al. 2004), or instead isthe sister taxon to Allosauroidea within a reconsti-
tuted Carnosauria (Currie 1995; Rauhut 2003) (Figs 3,4).
The placement of many individual taxa within any ofthese frameworks also varies. For example, allosauroids
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Smith et al., 2007
Coelop
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Benson et al., 2010
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NMV P186076
CERATOSAU
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Zupay
saurus
Coelop
hyso
idea
“dilo
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Xu et al., 2009
Figure 5. Results of previous phylogenetic analyses of Tetanurae, 2007–2010. Taxa recovered as non-coelurosaurian tetanurans in thepresent study are indicated in bold.
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The phylogeny of Tetanurae 219
reliably comprise Allosaurus, Acrocanthosaurus, Neove-nator, Sinraptoridae and Carcharodontosauridae, but thereis disagreement about whether Neovenator and Acrocan-thosaurus are allied with allosaurids (Currie & Carpenter2000; Holtz 2000; Allain 2002; Novas et al. 2005; Smithet 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 beenplaced within Allosauroidea (Bonaparte 1986; Serenoet al. 1994) but each exhibits morphological evidence ofother affinities (Zhao & Currie 1994; Rauhut 2003; Holtzet al. 2004; Smith et al. 2007; Benson 2010a; Brusatteet al. 2010a; Zhao et al. 2010).
‘Megalosaurs’ pose an even greater and more complexproblem. Many of the taxa that have at one time beenreferred to Megalosauridae have now been dispersedelsewhere, but a large number of putative megalosaurspecies remain. These include forms that have also beencalled torvosaurids (Kurzanov 1989; Sereno et al. 1994,1996), streptospondylids (Kurzanov 1989) and eustrep-tospondylids/eustreptospondylines (Paul 1988a; Bakkeret al. 1992; Yates 2005), but which are now consideredmembers of Spinosauroidea (Megalosauroidea). Mega-losaurus has been suggested to belong here (Britt 1991)but this was only recently supported analytically (Benson,2010a). Similarities between Torvosaurus, Edmarka andEustreptospondylus 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 withinthis 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 adiverse spinosauroid (or megalosauroid) clade, there is lessconsensus regarding the relationships between these taxa,resulting in little consistent support for anything otherthan a spinosaurid/torvosaurid (megalosaurid) dichotomy.Other taxa remain ambiguously placed. For example,Afrovenator has been allied with both allosauroids (Rauhut2003) and spinosauroids (Sereno et al. 1994; Allain 2002;Holtz et al. 2004; Smith et al. 2007; Benson, 2010a), andPoekilopleuron has been considered a spinosauroid (Allain2002; Allain & Chure 2002) and an allosauroid (Benson2010a; Benson et al. 2010). Many other taxa are quitefragmentary, and have either been excluded from mostprevious analyses or remained ambiguously resolved (e.g.Duriavenator, Magnosaurus, Marshosaurus) (Fig. 5).
In summary, although a great deal of progress has beenachieved in recent years (measured mainly by increasedconsensus), several points of uncertainty remain in tetanu-ran phylogeny and are therefore of primary interest here.These are: (1) whether spinosauroids (= megalosauroids)and allosauroids form a clade, or are serially arranged
outside Coelurosauria; (2) whether ‘megalosaurs’ form avalid clade and, if so, its membership; (3) placement of frag-mentary forms of potential geographic and temporal impor-tance; and (4) placement of relatively well known but prob-lematical forms (e.g. Cryolophosaurus, Marshosaurus,Monolophosaurus, Neovenator and Piatnitzkysaurus).
Resolving these and other phylogenetic problems withinTetanurae will allow a more reliable analysis of the evolu-tionary history, diversity and biogeography of this importanttheropod clade.
Problematical taxa
Herrerasauridae and EoraptorThe primitive taxon Eoraptor, along with the variouspurported constituents of the Herrerasauridae (often includ-ing Chindesaurus, Herrerasaurus (with junior synonymsFrenguellisaurus and Ischisaurus) and Staurikosaurus;Sereno 1999), have remained controversial since theiroriginal descriptions. Despite the availability of completematerials of both Eoraptor and Herrerasaurus, signifi-cant disagreement remains regarding their phylogeneticplacement. These forms have been hypothesized as basaltheropods (Sereno et al. 1993; Sereno 1999, 2007; Ezcurra& Novas 2007; Nesbitt et al. 2009), basal saurischians(Langer 2004; Irmis et al. 2007), basal dinosaurs anddinosauriformes (Fraser et al. 2002).
This paper does not address this question directly, insteadfocusing on the relationships of definitive theropod taxa. Wemerely assume that both Eoraptor and Herrerasaurus aremore primitive than all the ingroup taxa included in thisanalysis, and thus can be used to polarize character statesfor these forms. Whether Eoraptor and Herrerasaurus liewithin or outside Theropoda is not strictly relevant to theirutility in this regard.
Coelophysoids and ceratosaursThe taxonomic and systematic history of many of theseforms has been reviewed in detail elsewhere (Rauhut &Hungerbuhler 2000; Carrano & Sampson 2004, 2008;Tykoski & Rowe 2004) and will only be summarized here.
Coelophysoids are now understood to represent a basalradiation of theropods, globally distributed but confinedto the Late Triassic and Early Jurassic (Carrano & Samp-son 2004; Tykoski & Rowe 2004). Once linked as a sistergroup to ceratosaurs (Rowe & Gauthier 1990; Sereno 1999;Tykoski & Rowe 2004), most workers now support a morebasal placement for coelophysoids (Forster 1999; Sampsonet al. 2001; Carrano et al. 2002; Rauhut 2003; Smith et al.2007; Benson 2010a). Correspondingly, true ceratosaurs aremore closely related to tetanurans than to coelophysoids.Their record extends from the Early Jurassic through theLate Cretaceous (Carrano & Sampson 2008).
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Recent work has called the monophyly of Coelo-physoidea into question (Rauhut 2003; Smith et al. 2007;Nesbitt et al. 2009), but these results are controversial anddo not bear directly on the issues addressed in our study.
Megalosaurs and spinosaursMegalosaurus and its taxonomic descendants have beenthe most persistently problematic theropod group almostsince 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 awide array of species (in this case, 48) spanning nearly theentire Mesozoic and almost every continent. Unfortunately,the type species M. bucklandii was based on a series ofdisarticulated and largely disassociated bones from severalquarries around the village of Stonesfield, in the recur-rent ‘Stonesfield Slate’ facies of the Taynton LimestoneFormation (Bathonian, Middle Jurassic; Boneham & Wyatt1993). 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 justi-fied. As a result, it was impossible to clarify what clade, ifany, might be based on Megalosaurus. This problem hadattendant nomenclatural difficulties, because several higherlevel taxa (Megalosauridae, Megalosauroidea) are foundedon the genus Megalosaurus and would have priority overmore recent terms should that genus prove valid.
Spinosaurus and similar forms have likewise posedtaxonomic and phylogenetic problems since their initialdiscovery (Stromer 1915; Charig & Milner 1986). Theunusual nature of spinosaurid vertebrae and jaws ledto widely diverging opinions about their relationships.Spinosaurus was initially given its own family (Stromer1915; Huene 1926b), within which it was sometimes linkedto 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 discoveryof Baryonyx along with newer specimens of Spinosaurusled to debate concerning the potential relationships ofthese two forms (Charig & Milner 1986, 1990, 1997; Paul1988a; Buffetaut 1989, 1992; Buffetaut & Ouaja 2002).They are now considered to represent subfamilies withina 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; Benson2010a) have indicated that ‘megalosaurs’ and spinosauridsprobably form a larger clade. It remains to be determinedwhether other more fragmentary forms can be placed withinthis group, which has been previously defined on a rela-tively small number of synapomorphies (Sereno et al. 1994,1998).
Materials and methods
Outgroup relationshipsEoraptor and Herrerasaurus serve as outgroup taxa. Inchoosing multiple outgroups, we allow for greater reso-lution of character states at root nodes (Barriel & Tassy1998). These two taxa have been considered basal to allother theropods in all recent analyses, regardless of whetherthey 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 unitsThis 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. Thisincludes forms often referred to as megalosaurs, spinosaursand allosaurs, and represents the majority of basal tetanurantaxa. Several taxa whose placement had not previously beenaddressed, or which remained uncertain, were also included(e.g. Cryolophosaurus, Leshansaurus, Marshosaurus, Piat-nitzkysaurus, Shidaisaurus).
In order to explore whether any of these forms hadcloser affinities with other clades of basal theropods, weincluded multiple OTUs from two major non-tetanuranclades. The first, Coelophysoidea, is typically consideredthe most basal neotheropod clade and is here documented byDilophosaurus and two species of Coelophysis; its mono-phyly has recently been questioned (Nesbitt et al. 2009).The second, Ceratosauria, has recently been hypothesizedto occupy a more derived position than coelophysoids (e.g.Carrano & Sampson 2008), and is represented here byElaphrosaurus, Ceratosaurus, Masiakasaurus and Majun-gasaurus. These taxa were chosen both to represent ingroupcharacter states for these clades (without resorting to the useof suprageneric taxa), and to reflect known state variationswithin each group.
Ornitholestes, Proceratosaurus and Compsognathuswere selected to represent relatively completely knownbut less derived members of Coelurosauria (e.g. Holtz2000). That is, these taxa are not considered to be derivedornithomimosaurs, oviraptorosaurs, deinonychosaurs etc.,and therefore exhibit at least some primitive coelurosauriancharacter states.
In total, the ingroup consisted of 59 OTUs. Our goal inselecting these taxa was to effect a compromise betweenmaximal inclusiveness and productive analysis. Numerousbasal tetanuran taxa are so fragmentary and preserve sofew codable characters that their inclusion would swampthe matrix with missing data. However, we also deem itinsufficient to exclude taxa simply on the basis that theyare not complete. Such forms may preserve importantdata, including unique combinations of character states,which significantly impact the phylogenetic results. We
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also include several taxa that cannot be diagnosed basedon unique character states, but which we consider arelikely to represent valid species given their temporal and/orgeographic situation.
Ingroup taxa are listed below, including information onhypodigm, provenance, and diagnostic features. The diag-noses presented below combine original and (where cited)previously published observations. They are contextualizedwithin that clade for which most of the listed features areunique. For currently non-diagnosable taxa (sensu stricto),we include comments on the potential viability of the formin lieu of a traditional diagnosis. Specimens in bold wereexamined by the authors firsthand; the remainder were stud-ied via published photographs and drawings.
Institutional abbreviationsAM: Albany Museum, Grahamstown; AMNH: AmericanMuseum of Natural History, New York; AODF: AustralianAge of Dinosaurs Museum of Natural History, Winton; BP:Bernard Price Institute, Johannesburg; BSP: BayerischeStaatssammlung fur Palaontologie und Historische Geolo-gie, Munchen; BYU: Brigham Young University, Provo;CEPUNL: Centro de Estratigrafia e Paleobiologia daUniversidade Nova de Lisboa, Lisboa; CM: CarnegieMuseum of Natural History, Pittsburgh; CMP: CanteraMas de la Parreta collection, Museo de Valltorta, Tirig;CPP: Centro de Pesquisas Paleontologicas Llewellyn IvorPrice, Peiropolis; 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: FieldMuseum of Natural History, Chicago; FPDM: FukuiPrefectural Dinosaur Museum, Katsuyama; FSL: Facultedes Sciences, Universite Claude Bernard, Lyon; GA:Seccion de Geologıa, Sociedad de Ciencias Aranzadi, SanSebastian; GC: Goucher College, Baltimore; HASMG:Hastings Museum and Art Gallery, Hastings; IFPUB: Insti-tut fur Geologische Wissenschaften der FU Berlin, Fach-bereich Palaontologie, Berlin; IMGP: Institut und Museumfur Geologie und Palaontologie, Georg-August-Universitat,Gottingen; IPS/IPSN: Institut Paleontologic Dr. MiquelCrusafont, Sabadell; IWCMS: Museum of Isle of WightGeology, Sandown; IVPP: Institute of Palaeontology andPalaeoanthropology, Beijing; KMV: Kunming Munici-pal Museum, Kunming; KPE: Earth Science Department,Kyungpook National University, Daegu; KS: Kyeong-nam Science High School Museum, Kyeongnam; LDM-LCA: Lufeng Dinosaur Museum-Lufeng Chuanjie A’na,A’na; MACN: Museo Argentino de Ciencias Naturales‘Bernardino Rivadavia’, Buenos Aires; MB: HumboldtMuseum fur Naturkunde, Berlin; MCCM: Museo de lasCiencias de Castilla-La Mancha, Cuenca; MCF: MuseoMunicipal ‘Carmen Funes’, Plaza Huincul; MCZ: Museumof Comparative Zoology, Harvard University, Cambridge;
MG/MGSP: Museu Geologico, Lisbon; MH: Naturhis-torisches Museum, Basel; MIWG: ‘Dinosaur Isle’ Museumof Isle of Wight Geology, Sandown; ML: Museu da Lour-inha, Lourinha; MLP: Museo de La Plata, La Plata;MM: Universite 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: Museum National d’HistoireNaturelle, Paris; MNHNUL: Museu Nacional de HistoriaNatural da Universidade de Lisboa, Lisbon; MNN: MuseeNational du Niger, Niamey; MOAL: Molina Alto collec-tion, Museo de la Fundacion Conjunto Paleontologico deTeruel-Dinopolis, Teruel; MOR: Museum of the Rockies,Bozeman; MPCA: Museo Provincial ‘Carlos Ameghino’,Cipoletti; MPEF: Museo Paleontologico ‘Egidio Feruglio’,Trelew; MPM: Museo Padre Molina Paleontologıa deVertebrados, Rıo Gallegos; MPZ: Museo Paleontologicode la Universidad de Zaragoza, Zaragoza; MSNM: Museodelle Scienze Naturale di Milano, Milano; MUCP: Museode la Universidad Nacional del Comahue, Neuquen; MWC:Museum of Western Colorado, Fruita; NCSM: NorthCarolina 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: Paleontolog-ical Institute, Russian Academy of Sciences, Moscow;PS: Colectivo Arqueologico-Paleontologico de Salas, Salasde los Infantes; PVL: Fundacion Miguel Lillo, Univer-sidad Nacional de Tucuman, San Miguel de Tucuman;PW: Paleontological Collections, Department of MineralResources, Bangkok; QG: National Museum of NaturalHistory, Bulawayo; QW: Giant Buddha Temple Museum,Leshan; ROM: Royal Ontario Museum, Toronto; SDM:Stroud and District Museum, Dorset; SDSM: South DakotaSchool of Mines, Rapid City; SGM: Ministere de l’Energieet des Mines, Rabat; SM: Sirindhorn Museum of Palaeon-tology, Sahat Sakhan; SM∗: Senda Miravete collection,Museo de la Fundacion Conjunto Paleontologico de Teruel-Dinopolis, Teruel; SMA: Sauriermuseum Aathal, Aathal;SMNS: Staatliches Museum fur Naturkunde, Stuttgart;SMU: Southern Methodist University, Dallas; SNGM:Servicio Nacional de Geologia y Minero, Santiago deChile; TATE: Tate Museum, Casper College, Casper; TUB:Technishe Universitat Berlin, Berlin; UA: Departement dePaleontologie, Universite d’Antananarivo, Antananarivo;UCMP: University of California Museum of Paleon-tology, Berkeley; UC OBA: Department of OrganismalBiology 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) Universityof 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 Sao Paulo, Sao Paulo; UT: Geology Department, Univer-sity Al Fateh, Tripoli; WAM: Western Australia Museum,Perth; YNUGI: Geological Institute, Yokohama NationalUniversity, Yokohama; YPM: Peabody Museum of Natu-ral History, Yale University, New Haven; ZDM: ZigongDinosaur 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 incom-plete skull and partial postcranial skeleton (Stovall &Langston 1950).
Hypodigm. Holotype; OMNH 10147 (= MUO 8-0-S8), apartial skeleton (Stovall & Langston 1950); NCSM 14345,a nearly complete skull and skeleton (Currie & Carpenter2000); and SMU 74646, an incomplete skull and skeleton(Harris 1998a).
Diagnosis. Allosauroid theropod with: (1) absence ofnasal extension of antorbital fossa and associated pneu-matopores; (2) supraoccipital expanded parasagittallyinto double boss posterior to nuchal crest (Currie &Carpenter 2000); (3) cervical vertebral neural spines withtriangular anterior processes that insert into fossae ventral tooverhanging processes on preceding neural spines (Harris1998a); (4) neural spines of presacral, sacral and anteriorcaudal vertebrae more than 2.5 times taller than respectivecentrum lengths (Stovall & Langston 1950); and (5) acces-sory process on lateral surface of caudal prezygapophysis(Stovall & Langston 1950).
Occurrence. McLeod Prison, Arnold Farm and CochranFarm, Atoka and McCurtain Counties, Oklahoma, andHobson Ranch, Parker County, Texas, USA; Antlers andTwin Mountains Formations; late Aptian–early Albian,Early Cretaceous.
Remarks. Discovered more than half a century ago, Acro-canthosaurus has occupied many systematic positionsamong the large-bodied ‘carnosaurs’ since then, with vari-ous workers suggesting affinities with allosaurids (Stovall& Langston 1950; Colbert & Russell 1969), spinosaurids(Walker 1964; Romer 1966; Steel 1970; Carroll 1988) andmegalosaurids (Romer 1956; Swinton 1970; Russell 1984).More recently, consensus has favoured placement of Acro-canthosaurus within Allosauroidea, and debate has focusedon whether its closest ingroup affinities lie with Carchar-odontosauridae (Sereno et al. 1996; Harris 1998a; Sereno1999; Brusatte & Sereno 2008; Benson 2010a; Benson et al.
2010; Eddy & Clarke 2011) or Allosauridae (Holtz 1994a,2000; Currie & Carpenter 2000; Novas et al. 2005; Smithet al. 2007). Notably, the eponymous ‘high spined’ dorsalvertebrae have deep lateral embayments, resulting in an‘I-beam’ cross-section that is shared with Giganotosaurusand 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 morphol-ogy. Like Giganotosaurus and Carcharodontosaurus, theserecent finds have demonstrated the gigantic size of thistaxon, which approached the dimensions (although not themass, based on femoral circumference) of the largest tyran-nosaurids.
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 andskeleton.
Diagnosis. Allosauroid theropod with: (1) robust, cylin-drical transverse processes on proximal caudal vertebrae;and (2) fossa on lateral surface of coracoid dorsal to glenoidand (separate) subglenoid fossa.
Occurrence. Canadon Amarillo, Mendoza, Argentina;Anacleto Formation, Rıo Colorado Subgroup, NeuquenGroup; early Campanian, Late Cretaceous (Dingus et al.2000; Leanza et al. 2004).
Remarks. Sereno et al. (2008) were uncertain of the affini-ties of Aerosteon within Tetanurae. Consequently, theylisted several features in its diagnosis that can now beidentified in Neovenator (pneumatic canal within poste-rior dorsal transverse processes), Acrocanthosaurus (dorsalneural 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 cannotbe compared in closely related taxa. Based on recentcomparative work (Benson et al. 2010), our emended diag-nosis comprises two probable autapomorphies that areabsent in all other allosauroids, including a likely sistertaxon 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 incompleteskull and skeleton.
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Diagnosis. Megalosauroid theropod with: (1) rounded,lobate anterior maxillary margin of antorbital fossa (Serenoet al. 1994); (2) low, rectangular neural spine on third cervi-cal vertebra (Sereno et al. 1994); (3) broad lateral flangeon metacarpal I for articulation with metacarpal II (Serenoet al. 1994); and (4) posterior extension of distal pubicexpansion inset from lateral surfaces of anterior extension,forming pair of conjoined, posterior flanges.
Occurrence. In Abaka, Agadez, Niger; Tiouraren Forma-tion, Irhazer Group; Middle–Late Jurassic or Neocomian,Early Cretaceous (Lapparent 1960; Sereno et al. 2004;Rauhut & Lopez-Arbarello 2009).
Remarks. Afrovenator is represented by one of the mostcomplete theropod specimens known from Africa, includ-ing portions of all major skeletal regions. Originallydescribed as a basal torvosauroid (megalosauroid) (Serenoet al. 1994), it has also been recovered as an allosauroid(Rauhut 2003). Afrovenator exhibits a number of featuresthat seem to ally it with megalosaurs or other basal teta-nurans. In particular, the long, low skull shows little orna-mentation, and the pelvic girdle more closely resemblesthat of Marshosaurus and Piatnitzkysaurus than those ofallosauroids in the relative sizes of the iliac peduncles andmorphology of the distal pubis. The maxilla bears a longanterior ramus, a ‘kinked’ ascending ramus and a largeimperforate maxillary ‘fenestra’ resembling those of mega-losaurs 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 Allosaurusjimmadseni 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; UMNHVP 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,
9473, 9480, 9500, 9502, 9505, 9514, 9709, 10360, 10386,10779, 11031, 11463, 12231, 16584–16585; UUVP 1403,3304, 3894, 3981, 6000 (‘neotype’ = DINO 2560); USNM2323, 3304, 4734 (topotype, probably also includingUSNM 2315, holotype, Labrosaurus ferox), 4737, 7336,8257, 8302, 8335, 8367, 8405, 8423; YPM 1879?, 1890(holotype, Creosaurus atrox), 1931 (holotype, Labrosauruslucaris). A. jimmadseni: DINO 11541; MOR 693; SDSM30510; SMA 0005; UMNH VP C481.
Diagnosis. Allosauroid theropod with: (1) tall, mediolat-erally compressed dorsal projection (‘horn’) on posterodor-sal surface of lacrimal (Chure 2001a); (2) reduced externalmandibular fenestra; (3) strongly downturned paraoccipi-tal 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 areknown from dozens of sites in Utah, Arizona, NewMexico, Colorado, Wyoming, Montana and South Dakota,USA (cf. Turner & Peterson 1999; Foster 2003); SaltWash and Brushy Basin Members, Morrison Forma-tion; Kimmeridgian–Tithonian, Late Jurassic. A. europaeusderives from Andres, Praia de Vale Frades and Guimarota,Leiria, Portugal; Alcobaca and Porto Novo Members, Lour-inha Formation; late Kimmeridgian–Tithonian, Late Juras-sic (Perez-Moreno et al. 1999; Rauhut & Fechner 2005;Mateus et al. 2006).
Remarks. One of the most abundant large theropods in theworld, and consequently one of the best known, Allosaurushas been the subject of two monographic descriptions(Gilmore 1920; Madsen 1976a) and numerous studieson its growth and ontogeny (Bybee et al. 2006; Loewen2009), population variation (Chure & Madsen 1996; Smith1998), pathologies (Chure 2000; Hanna 2002) and taxon-omy (Smith 1998; Chure 2001a). In the last century ithas also largely occluded Megalosaurus as an archetypal‘basic’ theropod. Most of this material pertains to A. frag-ilis, although the recent discovery of a second North Ameri-can species has necessitated the transfer of some specimensinto the new taxon.
A. jimmadseni is based on a nearly complete skull andskeleton from Dinosaur National Monument (Chure 2000).It was mentioned in print (Chure et al. 2006) but has beendescribed in detail only in a dissertation (Chure 2001a)and so is technically a nomen nudum. A. jimmadseni isapproximately equivalent to the ‘creosaur-type allosaurid’described by Bakker (2000; which does not include thetype of Creosaurus). Because descriptive research on A.jimmadseni is ongoing (D. Chure pers. comm.; M. Loewenpers. comm.), we do not discuss its morphology in detailbut include it in the hypodigm of Allosaurus.
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A. europaeus was described on the basis of a partialskull (ML 415) by Mateus et al. (2006) that has alsobeen regarded as belonging to A. fragilis (Malafaia et al.2007); certainly it is more similar to that species than toA. jimmadseni. Other Allosaurus material from Portugal(MNHNUL/AND.001, a partial pelvis and hind limb) wasreferred to A. fragilis (Perez-Moreno et al. 1999) and A.europaeus (Mateus et al. 2006) but may not be determi-nate to the species level and is here regarded as Allosaurussp. A juvenile maxilla from Guimarota (IFPUB Gui Th 4)has also been assigned to Allosaurus sp. (Rauhut & Fech-ner 2005). Regardless of the validity of A. europaeus, weconsider these specimens to be correctly assigned to thegenus Allosaurus.
Angaturama limai Kellner & Campos, 1996
1996 Angaturama limai Kellner & Campos: 152, figs 2–4.
Holotype. USP GP/2T-5, the anteriormost portion of arostrum.
Diagnosis. Spinosaurid with midline ridge on dorsalsurface of conjoined premaxillae extending further ante-riorly than in Baryonyx and Suchomimus (modified fromKellner & Campos 1996). Angaturama cannot currently bedistinguished from Irritator because it is based on non-overlapping skull material.
Occurrence. Unspecified locality in the Araripe Basin,southern Ceara, Brazil; Romualdo Member, SantanaFormation; Albian, Early Cretaceous (Kellner & Campos1996).
Remarks. Angaturama possesses Spinosaurus-like dentalcharacters such as the absence of serrations on the carinae.It also shows the presence of a midline crest on the dorsalsurface of the premaxilla, similar to those of Baryonyx andSuchomimus. However, it differs from other spinosauridsin that no other taxon shows both features in combination.Angaturama is based on very fragmentary remains that mayrepresent 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 spec-ulative until the two holotypic specimens can be compareddirectly, and we retain the two taxa as distinct OTUs.
Australovenator wintonensis Hocknull et al., 2009
2009 Australovenator wintonensis Hocknull et al.: 24, figs20–37.
Holotype. AODF 604, a partial skeleton lacking most ofthe skull.
Diagnosis. Allosauroid with: (1) lateral groove along ulnarshaft (Hocknull et al. 2009); and (2) anterior bevelling ofproximal 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 (Hock-null et al. 2009).
Remarks. The recently described holotype of Australove-nator is the most complete theropod skeleton known fromAustralia. The original description was accompanied by aphylogenetic analysis that placed it within Allosauroidea,close to Neovenator but just outside Carcharodontosauri-dae. Indeed, although the skull is incomplete, the leftdentary is quite similar to that of Neovenator in its slen-der proportions and the arrangement of neurovascularforamina; these taxa also share extensively pneumatizedribs (among allosauroids). A number of other featureslisted in the original diagnosis of Australovenator can befound in other theropods, particularly Megaraptor, Chilan-taisaurus and Fukuiraptor, suggesting that all these taxamay 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 adorsal vertebra.
Diagnosis. Spinosaurid theropod with: (1) midline knobat posterior end of conjoined nasals terminating in cruci-ate process (Charig & Milner 1986); (2) subrectangularlacrimal horn (Sereno et al. 1998); and (3) peg-and-notcharticulation between scapula and coracoid (Sereno et al.1998).
Occurrence. Smokejacks Brickworks (Ockley brick pit),Walliswood, Ockley, near Dorking and Ewhurst Brick-works, Surrey, and south-west coast of the Isle of Wight,England; Cypridea clavata zone, Upper Weald Clayand Wessex Formations; Barremian–early Aptian, EarlyCretaceous (Charig & Milner 1997; Hutt & Newberry2004).
Remarks. The discovery of Baryonyx represented the firstreasonably complete theropod from the English Wealdenin more than a century and a half of collecting. It alsobecame the best known spinosaurid until the discovery ofSuchomimus (Sereno et al. 1998). The unusual and highlyderived nature of the skull and skeleton led to initial contro-versies surrounding its anatomy and relationships (Charig
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& Milner 1986, 1990, 1997; Paul 1988a; Sereno et al. 1998),but subsequent finds have supported its placement withinthe Spinosauridae (sometimes within its own subfamily,Baryonychinae; Sereno et al. 1998; Holtz et al. 2004;Benson 2010a) and have clarified the identities and detailsof several skull bones.
Genus Carcharodontosaurus Stromer, 1931
1931 Carcharodontosaurus Stromer: 19, pl. 1, figs 1–15.
Included species. C. saharicus (Deperet & Savornin,1925) Stromer, 1931 and C. iguidensis Brusatte & Sereno,2007.
Hypodigm. C. saharicus: BSP 1922 X46 (‘genoholo-type’; destroyed), a partial skull and skeleton, 1922 X45(‘Spinosaurus B’; destroyed), cervical vertebrae and pedalphalanges; 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) andIGU3–6, 11, teeth, skull elements, and a cervical centrum.
Diagnosis. Allosauroid theropod with: (1) large para-condylar and internal carotid artery pneumatic recesses(Brusatte & Sereno 2007); (2) funnel-shaped basisphenoidrecess (Brusatte & Sereno 2007); (3) reniform posteriorcentrum face in postaxial cervicals, twice as wide as tall(Sereno et al. 1996); and (4) deep ventral keel on postaxialcervicals that approaches depth of centrum (Sereno et al.1996).
Occurrence. C. saharicus is known from Timimoun,Algeria, the Baharıje Oasis, Egypt and the Kem Kem region,Morocco; ‘Continental Intercalaire,’ Baharıje, and KemKem beds, respectively; Albian?–Cenomanian, late Early-early Late Cretaceous (Stromer 1931; Lapparent 1960;Brusatte & Sereno 2007). C. iguidensis is known fromIguidi, west of In Abangarit, Agadez, Niger; Echkar Forma-tion, Tegama Group; Cenomanian, early Late Cretaceous(Brusatte & Sereno 2007).
Remarks. Like many large Jurassic and Cretaceoustheropods, Carcharodontosaurus was originally describedas a species of Megalosaurus (M. saharicus, Deperet &Savornin 1925) and has the rather unusual distinctionof having been referred to two different species of thistaxon (M. africanus, Huene 1956, p. 489 [incorrectlyattributed by that author to Deperet & Savornin 1925 andpossibly a lapsus calami]). In a later paper (Deperet &Savornin 1928), the original authors suggested an affili-ation with the North American genus Dryptosaurus, butsince Rauhut (1995) Carcharodontosaurus has uniformlybeen recognized as an allosauroid (e.g. Sereno et al.
1996; Brusatte & Sereno 2007). The syntype series ofM. saharicus consists of two teeth. However, despitehis reservations about the diagnostic utility of isolatedtheropod teeth, Stromer (1931, p. 10) specified this taxonas the type species of the new genus Carcharodontosaurus,designating a more complete specimen from Egypt asthe ‘genoholotype’ (Stromer 1931; reviewed by Rauhut1995).
The syntype teeth are now considered lost and Brusatte& Sereno (2007) designated a partial skull (SGM Din-1) from the upper Kem Kem beds, penecontemporane-ous with the Baharıje Beds in Egypt, as the neotype ofC. saharicus; it was claimed that “these Egyptian fossilswere never cast” (Brusatte & Sereno 2007, p. 904) butin fact an endocast from BSP 1922 X46 was made andcurrently resides in Berlin as MB.R.2056. The neotypeincludes much of the skull and indicates an individualof giant size. Using a reconstructed premaxilla based onthose of Giganotosaurus and Mapusaurus, we estimate theskull length of this specimen as close to 142 cm, equiva-lent to large specimens of Tyrannosaurus rex (e.g. FMNHPR 2081, at 139 cm). The femoral cross sectional propor-tions of BSP 1922 X46 suggests an animal of lower totalmass than a Tyrannosaurus of approximately equal femorallength.
Largely because of the distinctive crenulations presenton the teeth of Carcharodontosaurus, the genus has beenidentified across mid-Cretaceous deposits of North Africa,primarily from the ‘Continental Intercalaire’ and its descen-dant units; these beds vary in age from Albian throughCenomanian. The characteristic crenulations of isolated‘Carcharodontosaurus sp.’ teeth are present in other derivedcarcharodontosaurids such as Giganotosaurus (MUCPv-Ch 1) and Mapusaurus (Coria & Currie 2006), vary inprominence between individuals and along the tooth row,and are observed in a lesser developed form in numer-ous other tetanuran taxa (Brusatte et al. 2007). Giventhe recent identification of a second species from Niger(Brusatte & Sereno 2007), we consider it prudent torefer such isolated teeth to cf. Carcharodontosauridaeindet.
In addition, a complicated debate has surrounded theidentity of two fragmentarily known forms from NorthAfrica. The first, called ‘Spinosaurus B’ by Stromer (1934,p. 21), is represented by a vertebral series from the BaharıjeBeds of Egypt. The second specimen, based on severalisolated cervical vertebrae from the Kem Kem Beds ofMorocco, 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 thevertebrae of ‘Spinosaurus B’ and Sigilmassasaurus resem-bled one another, they were distinct from those associatedwith the Carcharodontosaurus vertebrae from BSP 1922
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X46. The issue warrants more extended treatment else-where, and at present we do not use specimens of Sigilmas-sasaurus for our character codings.
Ceratosaurus nasicornis Marsh, 1884
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 skulland partial skeleton.
Hypodigm. Holotype and MWC 1.1 (holotype,Ceratosaurus magnicornis), complete skull and partialskeleton; UMNH VP 5278 (holotype, Ceratosaurusdentisulcatus), 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, caudalvertebrae; 4951, 4952, 8907, 9142–9144, dorsal verte-brae; 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 (modi-fied from Marsh 1884); (2) medial oval groove on nasalsbehind 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, Canon City, FremontCounty, Fruita area, Mesa County and Dry Mesa Quarry,Montrose County, Colorado; Cleveland-Lloyd Quarry andSan Rafael Swell, Emery County, and Dinosaur NationalMonument, Uintah County, Utah; Quarry 9, Como Bluff,Albany County, Wyoming, USA; lower Brushy BasinMember, 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 considerthe existing North American specimens of Ceratosaurus torepresent 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,
and metatarsals; these likely pertain to the same individualas the lectotype (Hu 1964; Benson & Xu 2008).
Diagnosis. Tetanuran theropod with: (1) subrectangu-lar, anteromedially curving humeral deltopectoral crestprotruding almost as far anteriorly as it is long proximodis-tally, and bearing pitted scar on anterior surface (Rauhut2003; Benson & Xu 2008); and (2) obliquely oriented radial(lateral) condyle of humerus (Benson & Xu 2008).
Occurrence. Tashuikou (Dashuigou), 60 km north ofChilantai (Jilantai), eastern Alashan Desert, Nei MongolZizhiqu, China; Ulansuhai Formation; Turonian or younger(< 92 Ma), Late Cretaceous (Kobayashi & Lu 2003).
Remarks. This taxon was recently redescribed in detail,revealing several avetheropod synapomorphies (Benson &Xu 2008). Its taxonomic affinities remain uncertain butit is unlikely to belong to a megalosauroid (spinosauroid)as recovered by Rauhut (2003). Instead, Chilantaisaurusshares features with Fukuiraptor (long humerus relativeto femur; long deltopectoral crest; tall, narrow proximaldimensions of manual unguals), Aerosteon and Neovenator(preacetabular shelf on ilium) and Australovenator (narrowmanual unguals; median ridge on distal tibia) that suggest aclose 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; XiashaximiaoFormation; Middle Jurassic (He 1984).
Remarks. The holotype of Chuandongocoelurus shows apotentially diagnostic character combination as it possessestetanuran synapomorphies of the ilium (pubic peduncleof ilium considerably larger than ischial peduncle; pubicpeduncle approximately 1.34 times as long anteropos-teriorly as broad mediolaterally) and tibia (a vertical‘supraastragalar’ buttress located medially on the anteriorsurface of the distal end (Rauhut 2003) in combination witha femoral head that is directed approximately 45◦ antero-medially and inclined ventrally, otherwise seen only in non-tetanuran theropods. Chuandongocoelurus also possessesa hypertrophied supraacetabular crest, often considered anon-tetanuran feature (e.g. Holtz 2000), but this has alsobeen reported in Monolophosaurus (Zhao et al. 2010). Theilium and some dorsal centra are the only bones preservedin both taxa, and both show the same combination of teta-nuran and ‘non-tetanuran’ features. Therefore the two are
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only distinguishable in size: the Chuandongocoelurus holo-type represents an individual measuring 36 mm betweenthe pubic and ischial peduncles, whereas this measurementis 105 mm in Monolophosaurus (IVPP 84019).
He (1984) referred a second specimen (CCG 20011;vertebrae) to Chuandongocoelurus, but this materialrepresents a substantially larger individual and showslittle overlap with CCG 20010 that would allow positivereferral to the same taxon. Furthermore, CCG 20011possesses anteroposteriorly long cervical centra withanteroposteriorly elongate pleurocoelous fossa, similar tothose seen in the ceratosaur Elaphrosaurus. It is thereforeunlikely 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 skele-ton.
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; DMNH14729, 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, LateTriassic.
Remarks. Although C. bauri appears to be distinct from itsclose congener, C. rhodesiensis, the many available spec-imens of this taxon have not been subjected to detailedphylogenetic study for several decades. As a result, it canbe differentiated from other coelophysoids but not diag-nosed based on autapomorphies. We do not dispute thevalidity of C. bauri but note that further anatomical studyis still needed.
The type genus for Coelophysidae and therefore Coelo-physoidea, Coelophysis is a well-known small theropodrepresented by hundreds of individual specimens at the
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 agree-ment that the two taxa are extremely similar and probablyrepresent species of the same genus (Paul 1988a; Downs2000; 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 skullsand skeletons.
Diagnosis. Coelophysoid theropod with: (1) blunt, squaredanterior margin of antorbital fossa; (2) base of lacrimalvertical ramus width < 30% its height; and (3) maxil-lary and dentary tooth rows end posteriorly at ante-rior rim of lacrimal (all from Bristowe & Raath2004).
Occurrence. Maura River, Southcote farm and SpringGrange farm, Nyamandhlovu, Matabeleland North andChitake River, Mashonaland North, Zimbabwe; areabetween farms Edelweiss and Welbedacht, LadybrandDistrict, Free State, South Africa; Forest Sandstone andUpper Elliot Formations; ?Hettangian–Sinemurian, EarlyJurassic.
Remarks. Coelophysis rhodesiensis is one of themost abundantly preserved theropods, with hundredsof specimens representing dozens of individuals froma range of sizes and growth stages (Raath 1969,1977; Bristowe & Raath 2004). The material is in anexcellent state of preservation, with both articulatedand disarticulated individuals, making C. rhodesien-sis one of the most completely known early theropoddinosaurs.
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, Comp-sognathus corallestris), complete skull and skeleton; andMB.R.2003.2, cast of holotype counterslab.
Diagnosis. Coelurosaur with a possibly unique charactercombination 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 digitII; and (5) fourth trochanter on femur absent (all from Peyer2006).
Occurrence. Reidenburg-Kelheim area, Bayern, Germanyand near Bessons, Petit Plan de Canjuers, Var, Provence-Alpes-Cote d’Azur, France; Ober Solnhofen Plattenkalkand Canjuers Lithographic Limestone; Tithonian, LateJurassic (Bidar et al. 1972).
Remarks. Two nearly complete specimens document mostof the osteology of Compsognathus, and both have beenre-examined in recent years (Ostrom 1978; Peyer 2006). Allrecent phylogenetic analyses have agreed on its placementwithin Coelurosauria, although its specific relationshipshave been problematic. It is employed here as a relativelyunspecialized coelurosaurian, retaining some of the prim-itive features of the clade. Peyer (2006) provided a recentdiagnosis for C. longipes; although none of these featuresare autapomorphies, they may represent a unique charactercombination.
Concavenator corcovatus Ortega et al., 2010
2010 Concavenator corcovatus Ortega et al.: 203, figs 1,3–4.
Holotype. MCCM-LH 6666, nearly complete skull andskeleton with soft-tissue impressions.
Diagnosis. Allosauroid with: (1) four pneumatic recessesin nasal, three of which are connected; (2) rounded postor-bital brow occupying one-third of orbital fenestra; (3) dorsalvertebrae 11–12 with neural spines five times centrumheight; (4) caudal vertebrae 2–3 with tall, anteriorly angledneural spines; and (5) pollex terminates at or below distalend of phalanx I of digit II (all from Ortega et al. 2010).
Occurrence. Las Hoyas locality, La Cierva township,Cuenca, Spain; Calizas de La Huerguina Formation; lateBarremian, Early Cretaceous (Ortega et al. 2010).
Remarks. The remarkably well-preserved holotype indi-vidual of Concavenator was recently described as abasal carcharodontosaurian, approximately contemporane-ous with Neovenator salerii. Although its unusual vertebralmorphology superficially resembles that of Becklespinaxaltispinax (q.v.), Concavenator is quite distinct. UnlikeBecklespinax, the posterior dorsal centra bear large lateralfossae (although no foramina), the neural spines have lesscurved anterior and posterior margins at their bases, andthe apices of the tallest spines are anteroposteriorly narrow,curving towards a single apex formed from multiple spines.
The impression that Becklespinax possesses a combinationof tall and short posterior dorsal neural spines (Naish &Martill 2007) similar to those of Concavenator results froma broken anterior neural spine (confirmed by direct obser-vation of NHMUK R1828).
Condorraptor currumili Rauhut, 2005a
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 verte-brae, rib fragments, chevron, partial ilium, pubes, ischium,femora, metatarsal IV, and pedal phalanx (probably fromthe same individual as the holotype).
Diagnosis. Megalosauroid theropod with: (1) pleurocoel inanterior cervical vertebrae located immediately posterodor-sal to parapophysis; (2) shallow depression on lateralsurface of tibia at base of cnemial crest; and (3) metatarsalIV with distinct step dorsally between shaft and distal artic-ular facet (all from Rauhut 2005a).
Occurrence. Las Chacritas, 2.3 km west of Cerro Condor,Chubut, Argentina; Canadon Asfalto Formation, Sierra deOlte Group; Bajocian–Callovian, Middle Jurassic (Rauhut2005a; Volkheimer et al. 2008).
Remarks. This Middle Jurassic theropod, penecontempo-raneous with Piatnitzkysaurus, was discovered and diag-nosed recently (Rauhut 2005a). The fragmentary holotypepreserves elements from across the skeleton, providing themeans for comparison with other theropods. Its prove-nance makes it potentially important both evolutionarilyand biogeographically.
In addition to the holotype, Rauhut (2005a) referredother, disarticulated topotypic remains to Condorraptor onthe basis that they all represented a medium-sized theropodwith a similar quality of preservation, and thus could belongto a single individual. Most of these remains are closelycomparable with those of Piatnitzkysaurus, although theydo differ in some respects, suggesting that Condorrap-tor represents a related but distinct taxon. Rauhut (2005a)described the absence of a posterior incision between thefibular condyle and the medial part of the proximal tibiaas an additional autapomorphy. However, the proximal endof MPEF-PV 1672 is highly abraded and the absence ofthis incision is uncertain. In addition, the presence of aforamen on the lateral surface of the ischial peduncle ofthe ilium is present variably in other theropods, includ-ing Megalosaurus and Piatnitzkysaurus (Benson 2009b),and is not autapomorphic for Condorraptor (contra Rauhut2005a).
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Cryolophosaurus ellioti Hammer & Hickerson, 1994
1994 Cryolophosaurus ellioti Hammer & Hickerson: 828,figs 2, 3.
Holotype. FMNH PR 1821, posterior portion of skull withpartial postcranial skeleton.
Diagnosis. Theropod with: (1) transversely oriented,curved midline lacrimal crest bearing fluted anterior andposterior surfaces; (2) lower temporal fenestra constrictedby approximated jugal and squamosal; and (3) elongateanterior processes on cervical ribs (all from Smith et al.2007).
Occurrence. Mt. Kirkpatrick, near Beardmore Glacier,central Transantarctic Mountains, Antarctica; HansonFormation; Sinemurian–Pliensbachian, Early Jurassic(Hammer & Hickerson 1999; Smith et al. 2007).
Remarks. Cryolophosaurus is an important taxon forbasal theropod studies, as it is represented by a reasonablycomplete skull and skeleton from a poorly sampled region(Antarctica) during an early interval of theropod evolution.The skull is distinctive, bearing an apomorphic transversenasolacrimal crest and a nearly bipartite lower temporalfenestra. A recently published study (Smith et al. 2007)has described its morphology and relationships in detail,suggesting that Cryolophosaurus originated just basal to thesplit between Ceratosauria and Tetanurae. That study alsofound it to belong to a clade of ‘crested’ theropods morederived than coelophysoids, along with Dilophosaurus, ‘D.’sinensis (see below) and Dracovenator. Our observationssuggest that several putative similarities among these formsare over-split features that all relate to the presence of acranial crest, and that other skeletal features amongst themare less congruent. Recoding of the characters describingthe cranial crest by Brusatte et al. (2010a) resulted in fail-ure to resolve the ‘crested’ clade, but the affinities of mostcrested 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 lack-ing the skull.
Occurrence. Wujiaba Quarry, suburbs of Zigong city,Sichuan, China; lower part, Shangshaximiao Formation;Oxfordian–early Kimmeridgian, Late Jurassic (Dong et al.1983).
Remarks. This specimen was described in detail and hasbeen exhibited in the Municipal Museum of Chongqingfor decades (Dong et al. 1983). Its initial assignment toSzechuanosaurus campi is untenable because (1) the typematerials of S. campi (IVPP V.235, V.236, V.238, V.239)consist of teeth, which are non-diagnostic; and (2) no teethare 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 itexhibits a broad, expansive axial neural spine, quite similarto those of Sinraptor and Yangchuanosaurus. Other aspectsof the skeleton demonstrate tetanuran and allosauroid affini-ties. We retain this specimen as distinct from the type of‘S.’ zigongensis because we cannot identify autapomor-phies shared between them, and the latter derives from theunderlying Xiashaximiao Formation.
‘Dilophosaurus’ sinensis Hu, 1993
1993 Dilophosaurus sinensis Hu: 65, fig. 1.
Holotype. KMV 8701, a nearly complete skull and skele-ton.
Hypodigm. Holotype and LDM-LCA 10, a nearlycomplete skull and skeleton.
Diagnosis. Theropod with vertical groove or channel onlateral 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 theropodsknown from the lower Lufeng Formation (Hettangian–Sinemurian), ‘D.’ sinensis has been only preliminarilydescribed (Hu 1993). Although it was originally referred tothe genus Dilophosaurus based on the presence of a promi-nent nasolacrimal crest (Hu 1993), subsequent authors havequestioned this assignment (Lamanna et al. 1998). Specif-ically, several features of the holotype (KMV 8701) differfrom the conditions seen in Dilophosaurus, including thenumber of premaxillary teeth, the shape of the premaxillaand lower temporal fenestra, the extent of the maxillarytooth row and the morphology of the external mandibularfenestra. Indeed, as suggested by Lamanna et al. (1998),the currently understood distribution of cranial crests intheropods undermines their utility as indicators of system-atic affinity. Nonetheless, more recent work has supportedan affinity between ‘D.’ sinensis, Dilophosaurus and othercrested 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 isknown elsewhere only in Allosaurus and Neovenator, andthe premaxilla is reminiscent of the morphology of teta-nurans rather than coelophysoids. In addition, Hu (1993)described the cervicals as opisthocoelous and possessing asingle pleurocoel, the scapular blade as long and narrow,and the pubic peduncle of the ilium as larger than theischial peduncle; these characters are seen in tetanurans andceratosaurs but not coelophysoids. Finally, the distal ischiaare fused, as in Neovenator, Elaphrosaurus and Sinraptor.
The two specimens constituting the hypodigm of ‘D.’sinensis are often confused, thanks largely to the use ofLDM LCA 10 (or casts thereof) rather than the holotype innumerous exhibit displays. The two skulls have quite differ-ent proportions: KMV 8701 is shorter and taller, with amore abbreviate premaxilla, whereas LDM-LCA 10 is longand low, close to the proportions of D. wetherilli. In addi-tion, the dorsal crests in LDM-LCA 10 appear to end fartherposteriorly than those of KMV 8701. However, in spite ofthese differences there are no distinctions in the coded char-acters used in our analysis. The two specimens also sharean unusual ‘channel’ along the posterior premaxilla adja-cent to the maxillary contact, a vertical anterior border ofthe maxilla (distinct from the condition in Dilophosauruswetherilli), and a similarly shaped and placed promaxillaryfenestra. We suggest that the two specimens represent theproducts 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 skele-ton.
Hypodigm. Holotype, UCMP 37303, a partial skull andpostcranial skeleton, and UCMP 77270, a skull with partialskeleton.
Diagnosis. Theropod with: (1) thin, paired nasolacrimalcrests extending vertically from skull roof, each with finger-like 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, CoconinoCounty, Navajo Indian Reservation, Arizona, USA; SiltyFacies, 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 presumablyontogenetic stages. The nasolacrimal crest, once consid-ered unique among theropods, now appears to have beenapomorphic in shape but not presence, as similar struc-tures are also found in distantly related forms. Unspeci-fied minor differences between UCMP 77270 and the otherspecimens of D. wetherilli caused Welles (1984) to suggestthat it belonged to a separate taxon. We note the presenceof ‘fused’ maxillary interdental plates in UCMP 37303compared to the ‘unfused’ interdental plates of UCMP77270, and the presence of a ‘shelf-like’ trochanteric shelfof the femur in UCMP 77270 versus the mound-liketrochanteric region of UCMP 37302. These may representindividual rather than interspecific variations.
Tykoski (2005) referred a partial skeleton from GoldSpring, Arizona (TMM 43646) to D. wetherilli. However,direct examination reveals minor differences (e.g. tallermaxillary interdental places, presence of a pneumatic fossaon 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 witha fragmentary postcranial skeleton.
Diagnosis. Megalosauroid theropod with: (1) ventral notchin posterior outline of braincase between exoccipital-opisthotics and basioccipital (Rauhut 2004); and (2)absence of femoral distal extensor groove (Allain 2005a).
Occurrence. Near Conteville, Calvados, Basse-Normandie, France; Procerites progracilis Zone, Pierrede Caen, Calcaires de Caen; middle Bathonian, MiddleJurassic (Allain 2002).
Remarks. Dubreuillosaurus represents one of the mostimportant European theropod discoveries of the lastcentury, not only because it includes a well-preserved skull,but also because it derives from the Calcaires de Caen andis therefore approximately contemporaneous with Mega-losaurus and Poekilopleuron. As such, it has the potentialto illuminate the materials assigned to both taxa and clarifytheir phylogenetic relationships.
As noted by Allain (2002, 2005a), Dubreuillosaurusshares numerous synapomorphies with ‘megalosaurs’such as Eustreptospondylus, Afrovenator and Torvosaurus.Indeed, the maxilla of Dubreuillosaurus is very similarto that of Afrovenator. Dubreuillosaurus shows distinctautapomorphies amongst megalosaurs, but other featureslisted as diagnostic (Allain 2002, 2005a) are difficult toassess due to lack of comparative material from closely
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related taxa, or their more widespread presence in relatedforms.
The postcranium differs sufficiently from that of Poek-ilopleuron to warrant inclusion in a distinct genus (Allain2005a), but there is limited overlap in materials betweenthese two taxa. Unfortunately, the lack of skull material forPoekilopleuron 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 grooveon dorsal surface of jugal process of maxilla contain-ing numerous pneumatic foramina; and (2) array of smallforamina in ventral part of maxillary articular surface forpremaxilla (all from Benson 2008a).
Occurrence. Greenhill, Sherborne, Dorset, England;Garantiana garantiana Subzone, Parkinsonia parkinsoniZone, upper part of Inferior Oolite Group; late Bajocian,Middle Jurassic (Waldman 1974; Benson 2008a).
Remarks. The single known specimen of this taxonincludes just the anterior third of a skull, collected inthe 19th century (Owen 1883) but only fully preparedyears later (Waldman 1974; Benson 2008a). The originalspecific diagnosis (Waldman 1974, p. 325) relied only ontooth count and dental features to distinguish it from otherspecies of Megalosaurus; these features are now knownto occur more widely throughout Theropoda. The skullbones do reveal a few phylogenetically relevant features,comparable to those found in other taxa from the MiddleJurassic of Europe such as Dubreuillosaurus and Eustrep-tospondylus (Benson 2008a). These include the presenceof a distinct ‘kink’ in the maxillary ascending ramus, aparadental groove of the dentary that is wide anteriorly andan enlarged, subcircular third dentary alveolus.
Elaphrosaurus bambergi Janensch, 1920
1920 Elaphrosaurus bambergi Janensch: 225, figs 1–5.
Holotype. MB.R. (unnumbered), partial skeleton lackingthe skull, distal forelimbs, ribs, and distal caudals.
Hypodigm. Holotype, MB.R.1755, radius and 1756, distalischium.
Diagnosis. Ceratosaur with: (1) thin ventrolateral laminabordering posterior cervical pleurocoel ventrally; (2)strongly concave ventral border of cervical vertebrae, with
apex above mid-height of anterior articular face; (3) scapu-lar blade breadth exceeding height of vertebral column; and(4) extremely wide iliac brevis fossa with nearly horizontalbrevis shelf (all from Rauhut 2003).
Occurrence. RD, dd, and Dysalotosaurus quarries,Kindope, north of Tendaguru, Mtwara, Tanzania; Middleand ?Upper Dinosaur Members, Tendaguru Formation;late Kimmeridgian–?late Tithonian, Late Jurassic (Janen-sch 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 consistentlyidentified 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, skullelements and teeth.
Diagnosis. Allosauroid theropod with: (1) enlarged, subtri-angular, laterally exposed promaxillary fenestra largerin size than maxillary fenestra; (2) circular accessorypneumatic fenestra on posterodorsal ramus of maxilla;(3) dorsoventral expansion of antorbital fossa ventral topromaxillary and maxillary fenestrae; (4) postorbital browbearing finely textured ovoid swelling above posterodor-sal corner of orbit; (5) postorbital medial process withplate-shaped projection fitted to articular slot on frontal; (6)postorbital articulation for jugal includes narrow, laterallyfacing facet; (7) enlarged prefrontal lacking ventral process,with subquadrate exposure on dorsal skull roof and withinorbit; and (8) low protuberance on frontoparietal suture (allfrom Sereno & Brusatte 2008).
Occurrence. G88 and other sites along the Gadoufaouaoutcrop, Agadez, Niger; Elrhaz Formation, Tegama Group;Aptian–Albian?, Early Cretaceous (Sereno & Brusatte2008).
Remarks. Eocarcharia shares several features withcarcharodontosaurids but appears to represent a primitivemember of that clade. In particular, the maxilla is less heav-ily sculptured, the postorbital-squamosal suture is planarrather than helical, and the prefrontal remains unfused tothe 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 ofa juvenile individual.
Hypodigm. Holotype and OUMNH J.29775, a left ilium.
Diagnosis. Megalosauroid theropod with: (1) shallowlacrimal fenestra incorporating second, smaller foramen(Sadleir et al. 2008); (2) squamosal with hypertrophiedventrolateral flange obscuring posterodorsal corner of lowertemporal fenestra in lateral view (Rauhut 2003); (3) absenceof ventral midline ridge on posterior cervical and anteriordorsal vertebral centra (Sadleir et al. 2008); (4) markeddepression located anteriorly on ventral surface of tenthpresacral vertebra (Sadleir et al. 2008); (5) lateral wall ofiliac brevis fossa nearly horizontal, exposing medial wallof fossa along entire length in lateral view; and (6) pubicpeduncle of ilium as broad anteroposteriorly as mediolater-ally (modified from Benson 2009a).
Occurrence. Summertown brick pit, Wolvercot and Little-more, Oxfordshire, England; Peltoceras athleta Zone,Stewart Member, Oxford Clay Formation and CorallianFormation; late Callovian, Middle Jurassic (Huene 1926a;Walker 1964).
Remarks. The holotype specimen of Eustreptospondylusoxoniensis has had a lengthy history in theropod studies,as it long represented the most complete theropod skele-ton from Europe. Unfortunately, the taxonomic historyof the specimen has been equally long and rather convo-luted. Originally assigned to Megalosaurus (Woodward &Sherborn 1890), for many years it was considered the bestspecimen of Streptospondylus cuvieri Owen, 1842 (a nomendubium; see below) based on presumed similarities with thematerial now called Streptospondylus altdorfensis (Nopcsa1905, 1906; Huene 1926a; Swinton 1955). It was assignedto its own genus by Walker (1964).
Although the holotype of Eustreptospondylus is asubadult specimen, its completeness makes it importantin deciphering ‘megalosaur’ anatomy and relationships. Inaddition, much of the skull, including the braincase, ispreserved and shows similarities to those of Torvosaurusand Piatnitzkysaurus (Bakker et al. 1992; Sereno et al.1994; Rauhut 2004). Nopcsa’s (1906) lengthy descrip-tion was never translated into English and so was notwidely cited afterwards; only recently has the taxon beenredescribed in appropriate detail (Sadleir et al. 2008).
Rauhut (2003) synonymized Eustreptospondylus withMagnosaurus based on the shared presence of a trans-versely and dorsoventrally expanded anterior end of thedentary, 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, asdiscussed below (see Magnosaurus) there are also severaldifferences in the detailed morphology of preserved mate-rials (Sadleir et al. 2008; Benson 2010b). Notably also,Magnosaurus is represented by a more mature individualthat is approximately the same size as the juvenile holotypeof Eustreptospondylus. We do not consider them sufficientlysimilar to warrant synonymy and treat them as separate taxahere.
Fukuiraptor kitadaniensis Azuma & Currie, 2000
2000 Fukuiraptor kitadaniensis Azuma & Currie: 1737,figs 2–18.
Holotype. FPDM-V97122, associated cranial and appen-dicular bones (including V96082443, a left humerus).
Hypodigm. Holotype; FPDM-V9712229, fragmen-tary left maxilla, dentary, teeth, cervical vertebrae,dorsal neural arch, and coracoid; V97082553, lefthumerus; 97081115, V97082120, V980723005, righthumeri; V990410001, manual phalanx I-1; V980801141,V980815162, V9912141, manual unguals; V97080937,V98072302, V99090901, V98120001–0002, leftfemora; V97122BNA2, V97122BNA12, V970730003,V97081201, V97081330, V970813046, V970821039,V980805018, V9708102884, V980813017, V9812638,right femora; V97081317, V970814001, V970820060,right tibiae; V98082026, pedal phalanx III-2; and 42isolated teeth (FPDM various nos.; Currie & Azuma 2006).
Diagnosis. Avetheropod with: (1) proportionally long fore-arm (ulna : humerus ratio = 0.92); and (2) pubic peduncleof ilium approximately as broad anteroposteriorly as medi-olaterally (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 manyphylogenetic analyses since its original description, but bothanatomical descriptions (Azuma & Currie 2000; Currie& Azuma 2006) have placed it as a carnosaur (sensuRauhut, 2003) outside allosauroids and megalosauroids(spinosauroids). It is not clear whether such a positionresulted from the fragmentary nature of the available mate-rials or the mixed phylogenetic signal derived from them.Azuma & Currie (2000) suggested that long oblique ‘bloodgrooves’ (interdenticular sulci) on the teeth were an autapo-morphy of Fukuiraptor but they have since been reported inother theropods, including Megalosaurus (Benson 2009b),
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and can therefore be used as a phylogenetically informativecharacter (Benson 2010a).
More recently discovered taxa, including Australovena-tor (Hocknull et al. 2009) and Aerosteon (Sereno et al.2008), exhibit similar character combinations. For example,although Fukuiraptor appears to possess an autapomorphi-cally long forearm (relative to the arm), a proportionallylong forelimb (relative to the hind limb) is also seen inChilantaisaurus.
Giganotosaurus carolinii Coria & Salgado, 1995
1995 Giganotosaurus carolinii Coria & Salgado: 225, figs1, 2.
Holotype. MUCPv-Ch 1, partial skeleton.
Hypodigm. Holotype and MUCPv-52, 95, a dentary.
Diagnosis. Allosauroid with two pneumatic foramina onmedial surface of quadrate (Coria & Salgado 1995).
Occurrence. South and west of El Chocon, Lake EsquielRamos Mexia, Neuquen, Argentina; Candeleros Formation,Rıo Limay Subgroup, Neuquen Group; ?late Cenomanian,Late Cretaceous (Leanza et al. 2004; Corbella et al. 2004).
Remarks. Giganotosaurus has received a great deal ofattention because of its enormous size, comparable to thatof other carcharodontosaurids and in some ways rivallingthe largest tyrannosaurids. However, although the holotypeis relatively complete, a full description has not yet beenpublished and only the braincase has been documented indetail (Coria & Currie 2002). The original diagnosis ofGiganotosaurus (Coria & Salgado 1995) contains numer-ous features found in more recently discovered specimensbelonging to Acrocanthosaurus (Currie & Carpenter 2000)and Mapusaurus (Coria & Currie 2006), and which there-fore can no longer be considered diagnostic.
The reconstructed skull (Coria & Salgado 1995) includesa posteriorly oriented quadrate that outlines a trapezoidallower temporal fenestra. This unusual configuration isgenuine but several other skull contacts are not preserved,leading to ambiguity regarding its total length. We believethe original skull reconstruction is likely too long (153 cm),and as with Carcharodontosaurus (see above) we considerGiganotosaurus to have had a skull almost exactly compa-rable in length to that of Tyrannosaurus. Likewise, ourmeasurements 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 loweroverall body mass.
Irritator challengeri Martill et al., 1996
1996 Irritator challengeri Martill et al.: 5, figs 2–4.
Holotype. SMNS 58022, a nearly complete skull lackingthe tip of the rostrum.
Diagnosis. Spinosaurid theropod with nasals exhibitingprominent median bony crest that terminates posteriorlyin knob-like, somewhat dorsoventrally flattened projection(Sues et al. 2002).
Occurrence. Near Buxexe, 5 km south of Santana doCariri, Araripe Basin, southern Ceara, Brazil; RomualdoMember, Santana Formation; Albian, Early Cretaceous.
Remarks. Originally described as a maniraptoran thero-pod (Martill et al. 1996), a more recent redescription hasidentified Irritator as a spinosaurid (Sues et al. 2002).The slightly disarticulated, subadult holotype skull is themost complete known for any spinosaurid, and providesimportant information about skull morphology and propor-tions in this group. It also highlights the differencesbetween baryonychines such as Baryonyx and Suchomimus,and spinosaurines such as Irritator, Angaturama andSpinosaurus.
The presence of a broad lateral shelf on the surangu-lar was listed as an autapomorphy of Irritator (Sues et al.2002), but a similar morphology is also present in Bary-onyx. However, the combination of spinosaurine featureselsewhere in the skull of Irritator serves to distinguishit from baryonychines. As the surangular is unknown inSuchomimus, it cannot be determined whether this featureis homoplastic within Spinosauridae or a symplesiomorphywithin 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 postcranialskeleton.
Diagnosis. Tetanuran theropod with distinct ventral ridgeon all sacral vertebral centra (Li et al. 2009).
Occurrence. Xiaogu, Qianwei County, Sichuan, China;Shangshaximiao Formation; Late Jurassic.
Remarks. Leshansaurus was assigned to Sinraptoridae inits original description (Li et al. 2009). However, the brain-case bears distinct similarities to those of Piveteausaurusand Dubreuillosaurus whereas the maxilla resembles thoseof Afrovenator and Duriavenator. There are no clearsynapomorphies with members of Allosauroidea. There-fore 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 spursanterior to neural spines of even most proximal caudalvertebrae (modified from Mateus 1998); and (2) medialcondyle of tibia half the transverse width of fibular condyle.
Occurrence. Peralta, near Lourinha, Estremadura, Portu-gal; Sobral Formation; late Kimmeridgian–early Tithonian,Late Jurassic.
Remarks. Lourinhanosaurus was originally described asan allosauroid (Mateus 1998) and subsequently recoveredas a metriacanthosaurid by Benson (2010a) and Bensonet al. (2010). The skeleton is preserved in articulationbut many potentially informative elements are incompleteor poorly preserved, and little systematically informativedetail 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 postcranialskeleton and dentaries.
Diagnosis. Megalosauroid theropod with obliquelyoriented foramina located ventrally on lateral surfaceof dentary that are elongated to resemble grooves.Magnosaurus can also be distinguished from all othertemporally and geographically proximate theropods(Benson 2010b).
Occurrence. Nethercomb, 1.6 km north of Sherbourne,Dorset, England; Stephanoceras humphriesianum Zone andSubzone, middle part of Inferior Oolite; early Bajocian,Middle Jurassic (Huene 1923, 1926a; Waldman 1974).
Remarks. Originally described as a species of Mega-losaurus and later allocated its own genus, Magnosaurus,this taxon is based on fragmentary remains that have provendifficult to interpret phylogenetically. Waldman (1974)focused on the morphology of the dentaries and asso-ciated teeth to support its assignment to Megalosaurusbut made few comparisons with other theropods. Rauhut(2003) found several potential synapomorphies of thedentary to support a relationship with Eustreptospondy-lus. In both cases, affinities with traditional ‘megalosaurs’were implied, but this was only demonstrated in thecontext of a cladistic analysis recently (Benson 2010a). Adetailed redescription of the holotype, including fragmen-
tary postcranial material, has also been published (Benson2010b).
Majungasaurus crenatissimus (Deperet, 1896)
1896 Megalosaurus crenatissimus Deperet: 188, pl. 4, figs4–8.
1928 Dryptosaurus crenatissimus (Deperet); Deperet &Savornin: 262.
1955 Majungasaurus crenatissimus (Deperet); 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, and92.343 (syntype series, Megalosaurus crenatissimusDeperet, 1896); MNHN MAJ-4 (holotype, Majungatho-lus atopus Sues & Taquet, 1979); FMNH PR 2008, 2100;and UA 8678.
Diagnosis. Abelisaurid with: (1) thickened, fused, highlypneumatic nasals bearing large, bilateral foramina; (2) thinnasal lamina separating left and right premaxillary nasalprocesses; (3) maxilla with 17 teeth; (4) frontals withmedian hornlike projection; and (5) pronounced medianfossa on sagittal crest (modified from Sampson et al. 1998;Krause et al. 2007; Sampson & Witmer 2007).
Occurrence. Meravana and Berivotra, MahajangaBasin, Mahajanga, Madagascar; Anembalemba Member,Maevarano Formation; Maastrichtian, Late Cretaceous(Deperet 1896; Thevenin 1907; Sampson et al. 1998).
Remarks. The history of this taxon, and of its juniorsynonym Majungatholus atopus, has been given in detailelsewhere (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 mostelements of the skull and postcranium.
Occurrence. Canadon del Gato, Cortaderas area, 20 kmsouth-west of Plaza Huincul, Neuquen, Argentina; Huin-cul Formation, Rıo Limay Subgroup, Neuquen Group;Turonian–Santonian, Late Cretaceous (Dingus et al. 2000;Corbella et al. 2004; Leanza et al. 2004).
Remarks. The third carcharodontosaurid described fromArgentina, Mapusaurus is known from a bonebed depositcontaining several disarticulated individuals, some of which
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approximated Giganotosaurus in size. Coria & Currie(2006) united Mapusaurus and Giganotosaurus as sistertaxa within the subfamily Giganotosaurinae.
It is difficult to reliably distinguish elements ofMapusaurus from their counterparts in Giganotosaurusindependent of their stratigraphical origin. Several of theautapomorphies provided for Mapusaurus (Coria & Currie2006) are not observable in Giganotosaurus, while othersmust be reinterpreted. It appears to differ from Gigan-otosaurus in lacking a second pneumatic foramen on themedial quadrate and in details of the topology of the nasalrugosities.
We cannot confirm any of the autapomorphies ofMapusaurus proposed by Coria & Currie (2006). Severalcannot be observed in Giganotosaurus due to incompletepreservation, including the presence of an upper quadrato-jugal process of jugal split into two prongs; a small anteriormylohyoid foramen positioned above dentary contact withsplenial; and a humerus with broad distal end and littleseparation between condyles (Coria & Currie 2006). Thesupposed fusion between metacarpals II and III is basedon a specimen (PVPH 108-48) that we consider to be adistal humerus. As figured (Coria & Currie 2006, fig. 26),the ilium of Mapusaurus is long and low relative to thatof Giganotosaurus. However, this bone is poorly preservedand its outer margins represent broken surfaces, so the orig-inal 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 (= UUVP3266), right premaxilla; 7824 (= UUVP 1846), rightmaxilla; (= UUVP 1864), left maxilla; 7825 (= UUVP4695), right maxilla; 6364 (= UUVP 40-555), left dentary;6367 (= UUVP 3454), left dentary; 6368 (= UUVP 3502),right dentary; 6372 (= UUVP 1845), left ilium; (= UUVP1182), right ilium; 6374 (= UUVP 2742), right ilium; 6384(= UUVP 40-295), left pubis; 6386 (= UUVP 1867), leftpubis; 6387 (= UUVP 4736), right pubis; 6379 (= UUVP2832), 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 promi-nent ventral keels on posterior cervical and anterior dorsalvertebrae; and (2) articular surface of pubic peduncledivided into anterior bulge and posterior concavity (modi-fied from Madsen 1976b).
Occurrence. Cleveland-Lloyd Dinosaur Quarry, EmeryCounty and Dinosaur National Monument, Uintah County,
Utah; DMNH loc. 882, Moffat County and Dry MesaQuarry, Montrose County, Colorado, USA; Brushy BasinMember, Morrison Formation; Kimmeridgian, LateJurassic.
Remarks. Marshosaurus was originally described fromdissociated remains found at the Cleveland-Lloyd Quarryin Utah (Madsen 1976b), a few years after another enig-matic theropod, Stokesosaurus, was also identified from thesame deposit (Madsen 1974). For years thereafter, isolatedmedium-sized theropod bones from the Morrison Forma-tion could only be assigned preliminarily to one or the othertaxon (e.g. Britt 1991). Recent discoveries have improvedthis situation. A new specimen of Marshosaurus fromDinosaur National Monument (CMNH 21704) providesconfirmation of the original associations of cranial mate-rials proposed by Madsen (1976b). A partial skeleton ofStokesosaurus was also recently reported from England(Benson 2008b) that permits postcrania to be more reliablyassigned. However, we must still base some associationson the knowledge that Marshosaurus is a relatively primi-tive tetanuran whereas Stokesosaurus is a tyrannosauroid.These can only be confirmed by the discovery of a partialskeleton that includes an ilium similar to the holotype.
Although similar to both Condorraptor and Piat-nitzkysaurus, Marshosaurus differs from both in exhibitinggently convex (rather than flat) anterior centrum faces inthe anterior and middle cervicals; this is likely plesiomor-phic. Madsen (1976b, fig. 3A) described an imperforatemaxillary fenestra divided into two pockets instead of theusual single pocket as an autapomorphy of M. bicentes-imus based on a referred left maxilla (UMNH VP 7825).However, this feature is difficult to identify in recentlyreferred materials (CMNH 21704) and may vary betweenindividuals. The pubis (UMNH VP 6387) shows a smallanterior bulge, absent in other megalosauroids, that formsa blunt ‘peg-and-socket’ articulation with the ilium. This isthe reverse of the condition seen in ceratosaurs, in whichthe ilium bears a ‘peg’ that fits into a ‘socket’ on the pubis(Carrano et al. 2002). Marshosaurus is more completelyknown than either Condorraptor or Piatnitzkysaurus andincludes better-preserved maxillae and braincases that sharederived 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 specimenslisted by Carrano et al. (2011).
Diagnosis. Abelisauroid with: (1) four anteriormostdentary teeth procumbent, with first set in large, ventrallyexpanded alveolus that is almost horizontally oriented;
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and (2) strongly heterodont lower dentition, grading fromelongate, weakly serrated, apically round teeth with labi-olingually positioned carinae (anteriorly) to increasinglyrecurved, transversely compressed teeth with mesiodistallypositioned carinae (posteriorly) (all from Sampson et al.2001).
Occurrence. Berivotra, Mahajanga Basin, Mahajanga,Madagascar; Anembalemba Member, Maevarano Forma-tion; Maastrichtian, Late Cretaceous (Carrano et al. 2002).
Remarks. This taxon has been discussed in detail else-where (Carrano et al. 2002, 2011), including new materi-als that clarify previously unknown aspects of its anatomy(Carrano et al. 2011). More than two-thirds of the skeletonis 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 speci-mens listed by Benson (2010a).
Diagnosis. Megalosauroid theropod with a unique combi-nation of: (1) 13–14 dentary teeth (modified from Wald-man 1974); (2) unexpanded third dentary alveolus (Benson2010a); (3) dentary straight in dorsal view with unex-panded symphyseal area (modified from Waldman 1974);(4) lateral row of neurovascular foramina housed inshallow longitudinal groove in lateral dentary surface(Benson 2010a); (5) tall, unfused paradental plates ondentary (Benson 2010a); (6) two Meckelian foramina(Benson 2010a); and (7) shallow Meckelian groove (Benson2010a).
The referred materials of M. bucklandii exhibit additionalautapomorphies, including: (1) evenly rounded ventralsurfaces of sacral centra 1 and 3–5, with angular, longitu-dinal ridge on ventral surface of sacral centrum 2 (Benson2010a); (2) dorsally directed flange at mid-height of scapu-lar blade (Waldman, 1974); (3) array of posterodorsallyinclined grooves on lateral surface of median iliac ridge(Benson 2010a); (4) anteroposteriorly thick ischial apronwith nearly flat medial surface (Benson 2010a); (5) promi-nent, rugose distal ischial tubercle (Benson 2010a); and(6) complementary groove-and-ridge structures on articu-lar surfaces between metatarsals II and III (Benson 2010a).
Finally, M. bucklandii possesses several characters thatare not unique but are absent in all other megalosaurids,such as: (1) pneumatic jugal; (2) unexpanded third dentaryalveolus; (3) dorsal neural spines more than 1.9 timescentrum height; and (4) anterolaterally inclined deltopec-toral crest. Uniquely among megalosauroids, the maxil-lary anterior process is taller dorsoventrally than it is longanteroposteriorly (Benson 2010a).
Occurrence. Stonesfield, near Woodstock, 19 km north-west of Oxford, and Sarsgrove and Workhouse Quarry,Chipping Norton, Oxfordshire; New Park Quarry andOakham Quarry, Gloucestershire, England; StonesfieldSlate, Taynton Limestone Formation, Chipping NortonLimestone Formation and Sharp’s Hill Formation; lowestmiddle Bathonian, Middle Jurassic.
Remarks. The first dinosaur formally described (Buckland1824), M. bucklandii has remained problematic ever since.Known from disassociated, and probably isolated, bonesdeposited allochthonously in the marine Stonesfield Slatefacies of the Taynton Limestone Formation, the type seriesof M. bucklandii was long suspected to comprise at leasttwo distinct theropod taxa (Allain & Chure 2002; Day &Barrett 2004). Without definitive associations of elements,it has been difficult to determine which materials belong toeach ‘taxon’.
In addition, for decades it was commonplace to ascribe tothe genus Megalosaurus theropod specimens derived fromLate Triassic through Late Cretaceous strata worldwide,resulting in at least 50 nominal taxa (Benson 2009a). Theconsequence of this was a diluted perception of the truemorphology of the original materials, and the emergence ofwidespread opinion that Megalosaurus was a truly gener-alized theropod in almost all respects. In subsequent yearsnew genera were named for most of the diagnostic speciesreferred to Megalosaurus, but only recently has M. buck-landii been adequately restudied.
This is unfortunate because aside from the histori-cal interest in M. bucklandii, many of the StonesfieldSlate theropod specimens resemble those of Torvosaurus(Galton & Jensen 1979; Britt 1991) and Eustreptospondy-lus (Bakker et al. 1992) and therefore may represent anearly member of their lineage. These materials suggest thatat least one Stonesfield Slate taxon was not ‘generalized’but exhibited a morphology characteristic of one particulartheropod clade.
New research has clarified these materials and themorphology of M. bucklandii (Benson et al. 2008; Benson2009b, 2010a). As a result, we can be much more confi-dent that the majority (perhaps all) large theropod speci-mens 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 ofthe genus Megalosaurus. Several autapomorphies of the
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topotype material allow referral of specimens from otherBritish Bathonian localities (Benson 2010a).
Megaraptor namunhuaiquii Novas, 1998
1998 Megaraptor namunhuaiquii Novas: 4, figs 1–3.
Holotype. MCF-PVPH 79, a right ulna, left manualphalanges I-1 and I-2, and distal right metatarsal III.
Hypodigm. Type and MUCPv 341, a partial postcranialskeleton.
Diagnosis. Tetanuran with: (1) blade-like olecranonprocess of ulna (Novas 1998); (2) manual phalanx I-1 subtri-angular in proximal view with dorsal portion wider thanventral (Novas 1998); and (3) distal end of metatarsal IVshaft narrower than midshaft (Calvo et al. 2004).
Occurrence. Sierra de Portezuelo and Futalognko Quarry,Lago Barreales, Neuquen, Argentina; Portezuelo Forma-tion, Neuquen 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 mate-rials that initially suggested coelurosaurian affinities (Novas1998). More recent discoveries in the Futalognko Quarryhave provided additional anatomical information that clar-ifies its phylogenetic position (Calvo et al. 2002, 2004;Porfiri & Calvo 2003). The enlarged ungual, originallyascribed to the pes, has been shown to belong to manualdigit I. In addition, the remainder of the skeleton showssimilarities to carcharodontosaurians and other basal teta-nurans, rather than coelurosaurs.
Calvo et al. (2004) noted that carcharodontosaurid teethhave been found in the same quarry as the MUCPv 341 butthat in the absence of articulated specimens they couldnot be referred unequivocally to Megaraptor. However,one of the revised diagnostic characters of Megaraptor, anelongate anterior pleurocoel in the mid-cervical vertebrae(Calvo et al. 2004, p. 567), is also present in Giganoto-saurus, Neovenator, some specimens of Allosaurus, and amid-cervical vertebra from Morocco assigned to Carchar-odontosaurus saharicus (UCPC OT6). Similarly, anotherputative autapomorphy, prominent anterior and posteriorcentrodiapophyseal laminae on proximal caudal vertebrae,can also be observed in Aerosteon and Orkoraptor.
Metriacanthosaurus parkeri (Huene, 1923)
1923 Megalosaurus parkeri Huene: 453.1964 Metriacanthosaurus parkeri (Huene); Walker: 109,
fig. 16.
Holotype. OUMNH J.12144, a partial postcranial skele-ton.
Diagnosis. Allosauroid with ventral surfaces of posteriordorsal centra flat and with breadth approximately 2/3 poste-rior height of centrum (Benson 2009a).
Occurrence. Jordan’s Cliff, Overcombe, 1.5 km north ofWeymouth, 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 Mega-losaurus, Metriacanthosaurus was removed to its owngenus by Walker (1964), who noted the unusual shape ofthe dorsal margin of the ilium, moderate elongation of thedorsal vertebral neural spines, and fused distal ischia andpubes. Unfortunately, the dorsal margin of the ilium is suffi-ciently damaged and incomplete that the outline proposedby Walker (1964, fig. 16d) cannot be confirmed and is hereregarded as doubtful. Nonetheless, the morphologies of theilium, ischium and dorsal vertebrae are distinct enough fromthose of Megalosaurus to warrant retention of Metriacan-thosaurus as a separate taxon. It also occurs considerablylater in time than M. bucklandii and shows various simi-larities with penecontemporaneous Chinese ‘sinraptorids’(Paul 1988a; Benson 2009a).
Monolophosaurus jiangi Zhao & Currie, 1994
1992 Monolophosaurus jiangjunmiaoi Dong.: 71 (nomennudum).
1994 Monolophosaurus jiangi Zhao & Currie: 2028, figs1–5.
Holotype. IVPP 84019, a complete skull and skeleton.
Diagnosis. Tetanuran with: (1) nasal process of premax-illa bifurcated posteriorly (Brusatte et al. 2010a); (2) lateralsurface of premaxilla with deep groove between subnarialforamen and foramen on base of nasal process (Brusatteet al. 2010a); (3) large midline crest formed by nasals withstraight dorsal margin nearly parallel to maxillary alveo-lar margin (modified from Zhao & Currie 1994); (4) twoenlarged, subequal pneumatic fenestrae in posterodorsalpart of narial fossa (Brusatte et al. 2010a); (5) lacrimal withdiscrete, tab-like process projecting dorsally above preor-bital bar (Brusatte et al. 2010a); and (6) rectangular frontals,much wider than long (width/length = 1.67) (Brusatte et al.2010a).
Occurrence. 34 km north-east of Jianjungmiao, JunggarBasin, Xinjiang Uygur Zizhiqu, China; lower Shishugou (=Wucaiwan) Formation; middle Bathonian–late Callovian,Middle Jurassic (Zhao & Currie 1994; Eberth in Brusatteet al. 2010a and Zhao et al. 2010).
Remarks. One of the few well-represented Middle Juras-sic theropods, Monolophosaurus was originally described
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as a megalosaurid, though this was explicitly considered asan evolutionary grade (Zhao & Currie 1994). Its distinc-tive skull has drawn much attention, but the holotype alsoincludes most of the axial column and some appendicularelements. Most phylogenetic analyses based on the origi-nal, brief description have recovered Monolophosaurus asan allosauroid (Sereno et al. 1996; Rauhut 2003; Holtz etal. 2004), with which it shares several pneumatic features ofthe skull. However, Smith et al. (2007) noted some compar-atively primitive features and recovered Monolophosaurusas a basal avetheropod, and Benson (2010a) found it to bea megalosauroid closely related to Chuandongocoelurus.Recent detailed descriptions of the holotype noted numer-ous unusual and apparently primitive features of the skele-ton, and the affinities of Monolophosaurus remain far fromcertain (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, partialskull and skeleton divided between the two institutions.
Hypodigm. Holotype; MIWG 5470, two vertebrae and6352, fragmentary postcranial skeleton.
Diagnosis. Allosauroid theropod with: (1) accessory inter-premaxillary peg-and-socket articulation in dorsal part ofpremaxillary symphysis (Brusatte et al. 2008); (2) largemaxillary fenestra approximately 1/6 length of maxillarytooth row (modified from Hutt et al. 1996); (3) transverseexpansion of anterior face of axial intercentrum (Brusatte etal. 2008); (4) lateral foramina on anterior surface of odon-toid (Brusatte et al. 2008); (5) ischial distal boot conjoinedanteriorly but divergent posterolaterally (Brusatte et al.2008); (6) femoral head both oriented anteromedially andinclined proximally (Brusatte et al. 2008); (7) robust ridgeon external surface of femoral lesser trochanter (Brusatteet al. 2008); (8) thumbprint-shaped depression on posteriorsurface of femoral shaft, lateral to proximal end of fourthtrochanter (Brusatte et al. 2008); (9) proximodistally short,notch-like extensor groove and nearly flat anterior surfaceof distal femur (Brusatte et al. 2008); and (10) subovalrugosity on medial surface of distal tibia (Brusatte et al.2008).
Relative to other allosauroids, Neovenator is autapomor-phic in possessing fusion of the cervical ribs to the posteriorcervical vertebrae. In fact, the presence of camellate inter-nal texture exposed externally on the parapophyseal facetsof the eighth and ninth cervical vertebrae may record thefusion process of these elements.
Occurrence. Cliffs near Grange Chine, south-west coastof Isle of Wight, England; Wessex Formation; lateHauterivian–early Barremian, Early Cretaceous.
Remarks. The holotype of Neovenator is the mostcompletely preserved theropod from the Wealden Groupof England. It was the first definite allosauroid recognizedfrom Europe (although others have since been described;Perez-Moreno et al. 1999; Rauhut & Fechner 2005; Mateuset al. 2006; Ortega et al. 2010) and has been consideredthe latest surviving allosaurid (Hutt et al. 1996; Smith etal. 2007). Since its original description, the placement ofNeovenator within Allosauroidea has varied, with someauthors (Holtz et al. 2004; Brusatte & Sereno 2008; Benson2010a; Benson et al. 2010) favouring a closer relationshipto Carcharodontosauridae than Allosauridae.
Recent restudy of Neovenator has revealed a suite ofdistinctive autapomorphies, along with numerous synapo-morphies of carcharodontosaurians (Naish et al. 2001;Brusatte et al. 2008). Many of the autapomorphies proposedby Brusatte et al. (2008) are also present in the recentlydescribed Aerosteon and Australovenator (Benson et al.2010), and others may also prove to be more widelydistributed 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 partialskeleton.
Diagnosis. Coelurosaurian with: (1) teeth of premaxillaprominent, larger than maxillary teeth and bearing flattenedapex; and (2) retroarticular process offset medially fromlateral 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 well-known coelurosaurs from the Jurassic. Although knownfrom a nearly complete skeleton (Osborn 1903, 1917),Ornitholestes apparently lacks many of the specializationsthat characterize most individual coelurosaur clades and asa result, like Compsognathus, it has held many phyloge-netic positions within Coelurosauria. For the purposes ofthis study, it is used as a representative coelurosaur becauseit 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.
Hypodigm. Holotype and MACN-CH 895, partial skulland skeleton.
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Diagnosis. Tetanuran with: (1) strongly inflated base ofmaxillary ascending process (Rauhut 2003); and (2) evenlyrounded ventral surfaces of most sacral centra, except sacral3 bears flat midline strip and sacral 5 is broad and flat(Benson 2010a).
Occurrence. Cerro Condor, 1.5 km west of the formerFarias store, Cerro Condor village, right bank of theriver, Chubut, Argentina; Canadon Asfalto Formation,Sierra de Olte Group; Bajocian–Callovian, Middle Jurassic(Bonaparte 1979; Volkheimer et al. 2008).
Remarks. Initially assigned to Allosauridae (Bonaparte1979, 1986; Sereno et al. 1996), Piatnitzkysaurus repre-sents a temporally and geographically important taxon.Today it remains one of the few well-represented theropodsfrom the Middle Jurassic and one of the only diagnos-able theropods from the Jurassic of South America. Thetwo known specimens were found in close associationalong with materials of the sauropods Volkheimeria andPatagosaurus (Bonaparte 1979, 1986).
Restudy of the hypodigm of Piatnitzkysaurus reveals thatit lacks all the proposed synapomorphies of the Allosauri-dae and Allosauroidea, instead presenting a set of featurescharacteristic of more basal tetanurans. For example, thepubis has an enclosed obturator foramen and both thepubis and ischium show only modest distal expansions. Themaxilla is quite similar to that of Marshosaurus in lackingan anterior ramus and bearing a curved ventral margin, aventrally extensive antorbital fossa, and two prominent rowsof foramina parallel to the tooth row on the lateral surface.
Although Piatnitzkysaurus is very similar to Condorrap-tor, it can be distinguished from this form on the basisof the sacral vertebral centra (in Condorraptor the secondsacral centrum is broad and flat, whereas the third is broadand gently concave mediolaterally; Rauhut 2005a). Like-wise, Marshosaurus has a straight dentary whereas that ofPiatnitzkysaurus 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 Piveteausaurusare insufficient to determine autapomorphies for thistaxon given that the braincase of Leshansaurus is sostrikingly similar. However, based on its provenance andmorphology the only reasonable candidate for synonymy
is Eustreptospondylus, from which it has been specificallydistinguished (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 orig-inally described as a specimen of Streptospondylus cuvieri(Piveteau 1923). Walker (1964) made it the type of a secondspecies of Eustreptospondylus, E. divesensis, but it was laterremoved to its own genus (Taquet & Welles 1977). Thebraincase is large and weathered, making identification ofcertain sutures difficult. For example, we could not confirmthe exact boundaries of the supraoccipital and exoccipi-tal near the foramen magnum, which were illustrated byTaquet & Welles (1977). Some workers have suggested thatPiveteausaurus might be synonymous with Megalosaurus(Buffetaut et al. 1991a; Buffetaut & Enos 1992), but the lackof braincase material from the latter taxon makes this specu-lative, especially given the stratigraphical distance betweenthese forms. Nonetheless, the relatively anteroposteriorlylong parietals and frontals are more similar to those of non-avetheropod tetanurans than allosauroids.
The absence of adult braincase specimens fromother large megalosauroids prevents determining whetherPiveteausaurus truly represents a distinct taxon. Thegeneral proportions are dorsoventrally tall and anteropos-teriorly narrow, resembling those of large bodied membersof other clades (e.g. Allosaurus; Madsen 1976a), but unlikethe comparatively long and low braincases of the juve-nile 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 onthe basioccipital apron ventral to the occipital condyleand more than two-thirds its width, as do spinosaurs andsome other megalosaurs (e.g. Dubreuillosaurus, Eustrep-tospondylus). It also exhibits laterally, rather than ventro-laterally, directed paraoccipital processes, resembling thecondition in megalosaurs such as Dubreuillosaurus andEustreptospondylus, 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 (Eudes-Deslongchamps); Huene: 451.
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Holotype. Musee du Caen collection (destroyed), a partialpostcranial skeleton including caudal vertebrae, ribs,gastralia, forelimb, and hind limb.
Hypodigm. Illustrations of type materials (Eudes-Deslongchamps 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 olecra-non 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, MiddleJurassic.
Remarks. The unfortunate demise of the holotype andonly known specimen of Poekilopleuron during World WarII has been amply documented (e.g. Bigot 1945; Allain &Chure 2002) and has contributed significantly to the ambi-guity surrounding the affinities of this taxon. The spec-imen originally comprised a partial postcranial skeletonwith several phylogenetically informative elements, suchas the astragalus, tibia, and forelimb. Although the circum-stances of its original discovery and acquisition (Eudes-Deslongchamps 1837) have lent doubt as to whether theelements were associated, the published details of theaccount suggest that most or all of the materials pertainedto a single individual.
Since its discovery, the geographical and geological prox-imity of Poekilopleuron with Megalosaurus bucklandii hasled many researchers to speculate that the two taxa mightbe synonymous, or that Poekilopleuron might represent adistinct species of Megalosaurus (formally renamed M.poikilopleuron by Huene (1923) to avoid homonymy withthe type species). A recent review of Poekilopleuron (Allain& Chure 2002) supported the separation of these taxa, butpartly because the authors considered M. bucklandii to bea nomen dubium. They tentatively assigned Poekilopleu-ron to the Spinosauroidea (i.e. Megalosauroidea), citing asingle synapomorphy (the relative length of the deltopec-toral crest) and the absence of other features claimed to betypical of more derived theropods.
However, it is not clear that all of these features areabsent in Poekilopleuron (Benson 2010a). For example, thedistalmost preserved chevrons, representing middle caudalelements, show a partial bend or kink, less than that seenin most allosauroids and coelurosaurs but more than inmany more primitive theropods (e.g. coelophysoids andceratosaurs). The long axis of the humeral distal condylesform an oblique angle with the longitudinal shaft axis,rather than a right angle, giving an ‘S’-shaped outline inlateral view unlike the condition in megalosaurids but simi-
lar to that in Allosaurus. The humerus also shows significantobliquity between the long axes of the proximal and distalcondyles, which can be observed despite the missing prox-imal head. Finally, the midshaft median tuberosity on theradial shaft resembles to a similar structure in Acrocan-thosaurus (NCSM 14345; Currie & Carpenter 2000).
The genus Poekilopleuron (frequently misspelt Poicilo-pleuron or Poikilopleuron) was often used in the 19thcentury to receive incomplete specimens of large theropods.These include Poicilopleuron minor Cope, 1878 (indeter-minate), Poikilopleuron pusillus Owen, 1876 (now Aris-tosuchus pusillus; Seeley, 1887), Poekilopleuron schmidtiKirprianow, 1883 (see Fragmentary Taxa, below) andPoicilopleuron valens Leidy, 1870 (now Antrodemusvalens; Leidy, 1873). None of these materials pertain toPoekilopleuron, which should be restricted to the typespecies.
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 mostof the dorsal surface.
Diagnosis. Coelurosaurian with: (1) anterior nasal processof premaxilla inclined slightly anterodorsally, with nasalhorn core overhanging premaxillary internarial bar anteri-orly; (2) premaxillary internarial bar bifurcated posteriorly;(3) promaxillary foramen located posterodorsal to anteriorend of antorbital fossa; and (4) anteriormost dentary toothcurved anteriorly, with carinae oriented labiolingually (allfrom Rauhut et al. 2010).
Occurrence. Minchinhampton Reservoir, Gloucester-shire, England; White Limestone Formation, Great OoliteGroup; middle–late Bathonian, Middle Jurassic.
Remarks. P. bradleyi is represented by most of a skullthat was originally described as a species of Mega-losaurus (Woodward 1910). Huene (1926a) later recog-nized its generic distinctiveness and highlighted the unusualstructure of the rostrum, which includes the base of amidline horn or crest; this led him to assign the taxonto Ceratosauria. More recently, Proceratosaurus has beenconsistently assigned to Coelurosauria (Holtz 2000; Rauhut2003; Holtz et al. 2004), and more specifically has beenconsidered a primitive tyrannosauroid (Rauhut et al. 2010).It is used here to represent basal coelurosaurian characterstates.
Saurophaganax maximus Stovall, 1941
1941 Saurophagus maximus Stovall in Ray: 38 (nomennudum).
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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 vertebralneural arch.
Hypodigm. Holotype and OMNH 01771, postorbital;01142, 02145, quadrates; 01152, 01153, 01680, teeth;01135, atlas; 01444, 01446, 02146, 02147, cervical verte-brae; 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 prezy-gapophyses for proatlas; and (2) accessory horizontallamina at base of dorsal vertebral neural spines, locateddorsal and parallel to transverse process (all from Chure1995).
Occurrence. Stovall Quarry 1, east of Kenton, CimarronCounty, Oklahoma, USA; upper Brushy Basin Memberequivalent, Morrison Formation; ?Tithonian, Late Jurassic(Stovall 1938; Chure 1995).
Remarks. The validity of Saurophaganax has beendebated since its discovery. Originally named Sauropha-gus maximus (Ray 1941; a nomen nudum: Chure 1995),the type materials derived from a bonebed that was inex-pertly excavated and prepared, resulting in poorly preservedmaterials for study (Chure 1995). The specimen is similarto Allosaurus and has been referred to that genus (Paul1988a; Smith 1998). However, Chure (1995) has identifiedat least two apomorphies on the skeleton. In addition, thelocality appears to derive from high within the MorrisonFormation, potentially above most other Allosaurus sites(Turner & Peterson 1999). We retain it here as a separatetaxon.
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.
Lectotype. IVPP V.2885.1–2885.2, a well-preservedbraincase and posterior part of the skull roof.
Hypodigm. Lectotype and paralectotype series: IVPPV.2885.3–V.2885.7, partial skull, axis and caudal vertebraethat likely pertain to the same individual as the lectotype(Hu 1964; Brusatte et al. 2009a).
Diagnosis. Allosauroid with: (1) maxillary antorbital fossareduced in extent, nearly absent; (2) paradental groove onmedial surface of maxilla absent; (3) deep, dorsoventrallyoriented grooves located dorsally on maxillary paradentalplates; (4) pneumatic recess penetrates to posterior end ofnasal; (5) sagittal crest on frontal; and (6) large pneumaticforamen at anterodorsal corner of dorsal tympanic recessof prootic (all from Brusatte et al. 2009a, 2010b).
Occurrence. Maortu, 60 km north of Chilantai (Jilantai),eastern Alashan Desert, Nei Mongol Zizhiqu, China; Ulan-suhai Formation; Turonian or younger (< 92 Ma), LateCretaceous (Kobayashi & Lu 2003).
Remarks. The second named species of Chilantaisaurus,C. maortuensis, is represented by a very partial skele-ton. The distinct morphologies of these species have ledto 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 havebeen found to belong to distinct genera (Chure 2001a),with ‘C.’ maortuensis placed in the new genus Shaochi-long 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.
Holotype. LDM-LCA 9701-IV, a partial skull and skele-ton.
Diagnosis. Allosauroid with supraoccipital excluded fromforamen magnum by midline contact between exoccipi-tals (otherwise unknown in tetanuran theropods; Wu et al.2009).
Occurrence. Quarry 1, Laochangjing, A’na village,Chuanjie township, Lufeng County, Yunnan, China; baseof Upper Lufeng Formation; Middle Jurassic (Wu et al.2009).
Remarks. The original description of Shidaisaurus (Wuet al. 2009) listed a unique combination of severalfeatures, including downturned paraoccipital processes, themorphology of the lamina between the axial neural spineand 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 sinraptoridsand other tetanurans and are best dealt with as descriptivecharacter states outside of the taxon diagnosis. In addi-tion, although Wu et al. (2009, p. 9) stated that pleu-rocoels were not present in “any anterior dorsal verte-brae” of Shidaisaurus, they also noted that none of thevertebrae prior to dorsal 4 are exposed. Given that mosttheropods express pneumaticity up to but not past dorsal4, the presence of this potential autapomorphy among non-coelurosaurian 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 Musee desDinosaures, Esperaza).
Diagnosis. Avetheropod with double vertical ridge oncentral part of lateral iliac blade (Rauhut 2003).
Occurrence. Phu Wiang 9, Amphoe Phu Wiang, Chang-wat Khon Kaen, Thailand; Sao Khua Formation;Barremian–Aptian, Early Cretaceous (Racey & Goodall2009).
Remarks. In their original description, Buffetaut et al.(1996) interpreted Siamotyrannus as an Early Cretaceoustyrannosaurid, noting particularly the development of verti-cal ridging on the lateral surface of the ilium. However,this feature is expressed to varying degrees of promi-nence among widespread basal tetanurans other thantyrannosauroids, including Allosaurus, ‘Iliosuchus’, Mega-losaurus and Piatnitzkysaurus (Bonaparte 1986; Benson2009b, 2010a).
The preserved elements of Siamotyrannus differ fromcorresponding tyrannosaur bones in several respects, espe-cially the pubis and ischium (Fig. 6). For example, the iliumis broken distally in Siamotyrannus, not naturally abbrevi-ated as in tyrannosaurs and other coelurosaurs. In fact thisbone is quite robust even up to the preserved break surface.The ischium further bears a midshaft bulge developed byposterior extension of the contacting medial surfaces of theischia, seen elsewhere in Sinraptor and Yangchuanosaurus.The pubic obturator foramen is open, but not broadly so asin tyrannosaurids, instead resembling the unusual conditionexhibited by Sinraptor and Yangchuanosaurus. Likewise,the pubic boot is oriented similarly to that in Sinraptor(strongly posterodorsally inclined). With a thick anterioredge and a narrower posterior portion, it is much closerto the condition seen in basal tetanurans than coelurosaurssuch as tyrannosauroids.
Figure 6. Left pelvic girdle of Siamotyrannus isanensis Buffetautet al., 1996 (Musee des Dinosaures cast of PW9-1), showingfeatures 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 tyran-nosauroids, where a single prominent ridge is typical. Theanterior edge lacks the bilobate shape seen in most tyran-nosaurids. Finally, although the pubic peduncle of the iliumis larger than the iliac peduncle, as is typical of tetanurans,the latter is not acuminate in Siamotyrannus, as would betypical 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 skele-ton.
Diagnosis. Allosauroid with: (1) enlarged lateral tempo-ral fenestra with relatively straight postorbital-squamosalbar; (2) very short squamosal ramus of postorbital; and (3)palatine very deeply pneumatic between internal naris andpostpalatine fenestra (all from Currie & Zhao 1994).
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Occurrence. Jianjungmiao, Junggar Basin, XinjiangUygur Zizhiqu, China; upper part, Shishugou Formation;Oxfordian, Late Jurassic.
Remarks. The well-preserved, complete skull and associ-ated partial skeleton of Sinraptor have been described andillustrated in detail, providing a keystone for the recogni-tion of an entire clade of theropods, Sinraptoridae (Currie& Zhao 1994). Although Sinraptor shares many featureswith allosauroids, it also retains a more primitive morphol-ogy in the manus and pelvis that resembles the condition inmegalosauroids.
Spinosaurus aegyptiacus Stromer, 1915
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 anteriorportion 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), fragmentaryjaws; and UCPC 2, a midline nasal crest.
Diagnosis. Spinosaurid with: (1) no midline crest onconjoined premaxillae (Dal Sasso et al. 2005); (2) premax-illa entirely excluded from borders of external naris (DalSasso et al. 2005); and (3) extremely elongate dorsal neuralspines (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 isbased on materials that survive only as lithographic plates(Stromer 1915), the originals having been destroyed inthe same World War II raid on Munich. However, unlikeCarcharodontosaurus, Spinosaurus did not enter into aperiod of obscurity thanks to the highly unusual nature ofthe type specimen. Characterized by enormously elongateneural spines, Spinosaurus and the family Spinosauridaehave always retained a place in both scientific and popularliterature.
This lasting fame has helped perpetuate the highly spec-ulative reconstruction made by Huene (1956, fig. 517)and the notion that Spinosaurus may have been partlyquadrupedal (Charig & Milner 1986). With little anatom-ical data available, most researchers either placed it alonewithin its own family (e.g. Stromer 1915; Huene 1926b;Piveteau 1955) or allied it with the only other long spinedgenera then known, Acrocanthosaurus and Altispinax (e.g.Walker 1964; Romer 1966; Steel 1970).
More recent discoveries across North Africa (Buffetaut1989, 1992; Buffetaut & Ouaja 2002; Russell 1996; DalSasso et al. 2005) have brought to light new materials ofSpinosaurus 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), butother workers have subsumed this within S. aegyptiacus(Sereno et al. 1998). We follow the latter course here, butnote that, as with Carcharodontosaurus, future discoveriesmay better support the presence of distinct species in thedifferently aged North African beds.
Streptospondylus altdorfensis von Meyer, 1832
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 indi-vidual.
Hypodigm. Holotype and MNHN 9645, the distal part ofa left femur.
Diagnosis. Tetanuran with bifurcated (or paired) hypa-pophyses on ventral surface of anterior dorsal vertebrae(Allain 2001; Benson 2010a, fig. 19B).
Occurrence. Calvados, Falaises des Vaches Noires, nearHonfleur, Basse-Normandie, France; Marnes de Dives; lateCallovian–early Oxfordian, late Middle–early Late Juras-sic.
Remarks. The partial remains of Streptospondylus weresome of the earliest dinosaur bones formally described,albeit intermixed with (and interpreted as) crocodylianremains at the time (Cuvier 1808; Allain 2001). Thematerial is very incomplete but comes from a poorlyknown time interval for theropod evolution and thus haspotential importance. Although the presence of bifurcatedhypapophyses has been used to unite Streptospondylusand Eustreptospondylus as sister taxa (Allain 2001), it isnot present in the latter form and is here considered anautapomorphy 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 posterolateralpremaxillary process that nearly excludes maxilla fromexternal naris; (2) posterior dorsal, sacral and anteriorcaudal vertebral neural spines expanded anteroposteriorlyand dorsoventrally; and (3) hook-shaped radial ectepi-condyle (all from Sereno et al. 1998).
Occurrence. Gadoufaoua outcrop, Agadez, Niger; ElrhazFormation, Tegama Group; Aptian–Albian?, Early Creta-ceous.
Remarks. Suchomimus is one of the better representedspinosaurids, with well-preserved material known from theskull, axial skeleton, and appendicular region of severalindividuals over a range of body sizes.
There has been discussion regarding the validity ofSuchomimus as a distinct genus from Baryonyx, with some(Sues et al. 2002) favouring referral of S. tenerensis to thelatter genus. Among the diagnostic features listed above forSuchomimus, character (1) cannot be observed in Baryonyxand (2) is difficult to assess because of the immature natureof the Baryonyx holotype. (A later study suggested that theneural spines over the pelvic region may also be expanded inBaryonyx [Charig & Milner 1990].) It remains subjectivewhether enlargement of the radial entepicondyle justifiesgeneric separation, but we retain the original designationhere. A fourth autapomorphy, presence of a hypertrophiedulnar olecranon process offset from the humeral articula-tion, was also listed originally (Sereno et al. 1998) butappears to be present in Baryonyx, although obscured byincomplete preservation of the proximal ulna.
‘Szechuanosaurus’ zigongensis Gao, 1993
1993 Szechuanosaurus zigongensis Gao: 308, figs 1–6, pls1–3.
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 2–4opisthocoelous, remainder amphiplatyan (Holtz et al.2004). Although the specimen requires detailed restudy,it can be distinguished from other theropods in the sameformation (Rauhut 2003).
Occurrence. Dashanpu Dinosaur Quarry, Zigong,Sichuan, China; Xiashaximiao Formation; Middle Jurassic(Gao 1993).
Remarks. The holotype skeleton of ‘S.’ zigongensis hasbeen included in a number of phylogenetic studies (Rauhut2003; Holtz et al. 2004), which have tended to recoverit among basal tetanurans. Unfortunately, the holotype ofSzechuanosaurus campi, the type species of the genus, isnon-diagnostic and therefore S. campi is a nomen dubium(Chure 2001a; Holtz et al. 2004). The species ‘S.’ zigongen-sis, however, is based on much of a postcranial skeleton thatChure (2001a) felt was sufficient to justify creation of a newtaxon to include it and CV 00214. We do not consider theseforms close enough to include in one taxon, and indeed theyderive from different strata and can be scored differently formany 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 (nomennudum).
Holotype. BYU-VP 2002, left and right forelimbs.
Hypodigm. Holotype and BYU-VP 2000 series; TATE401, 1002–1005 (holotype, Edmarka rex), jugal, scapulo-coracoid, and ribs (Bakker et al. 1992); 1003, 1012 (‘Bron-toraptor’), pelvis, femur? and other postcranial material(Siegwarth et al. 1997).
Diagnosis. Megalosauroid with: (1) very shallow maxil-lary fossa; (2) ‘fused’ paradental plates (Britt 1991); (3)pronounced rim around anterior face of cervical centra(Britt 1991); (4) expanded fossae in posterior dorsal andanterior caudal vertebrae centra forming enlarged, deepopenings (modified from Britt 1991) that we interpret aspneumatic; (5) highly ossified puboischiadic plate (modi-fied from Britt 1991); and (6) distal expansion of ischiumwith prominent lateral midline crest and oval outline inlateral view.
Occurrence. Dry Mesa Quarry, Montrose County; CalicoGulch Quarry, Uncompahgre Plateau, Moffit County, andMeyer site, Garden Park, north of Canon City, FremontCounty, Colorado; Carnegie Quarry, Dinosaur NationalMonument, Uintah County, Utah; Gilmore Quarry N,Freezeout Hills, Carbon County, and Nail and Louise Quar-ries, Como Bluff, Albany County, Wyoming, USA; SaltWash and Brushy Basin Members, Morrison Formation;Kimmeridgian–Tithonian, Late Jurassic.
Remarks. Torvosaurus was an important discovery fromthe Morrison Formation (Galton & Jensen 1979), not only
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because it represented a new large theropod, but alsobecause it appeared to be more primitive than Allosaurusand in many ways similar to Megalosaurus (Galton &Jensen 1979; Britt 1991). Later, additional specimens werediscovered at other Morrison Formation quarries. Amongthese, the incomplete materials that have been referredto Edmarka rex (Bakker et al. 1992) and ‘Brontoraptor’(Siegwarth et al. 1997) may represent species level vari-ants of Torvosaurus, but we do not consider the observeddifferences as sufficient to justify a new taxon at this time.Therefore we consider E. rex and ‘Brontoraptor’ as juniorsynonyms 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 toTorvosaurus sp. It is likely that this does indeed repre-sent Torvosaurus, although it may also belong to a relatedform; there are no clear autapomorphies on the tibia asidefrom its very robust proportions. Mateus et al. (2006)later referred a maxilla (ML 1100) from the base ofthe Lourinha Formation of Praia da Vermelha, Portu-gal to Torvosaurus tanneri. However, they noted a muchreduced tooth count of 10 (compared to 11–13 in the topo-type maxilla), which suggests a more conservative assign-ment to Torvosaurus sp. pending the discovery of furthermaterial.
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 skele-ton.
Diagnosis. Allosauroid with: (1) bilobate denticles onanterior carinae of teeth; and (2) very pronounced mentalgroove on lateral surface of dentary (all from Novas et al.2005).
Occurrence. Estancia ‘La Juanita’, 28 km north-east ofPaso de Indios, Chubut, Argentina; Cerro Castano Member,Cerro Barcino Formation; Aptian–Albian, Early Creta-ceous (Rauhut pers. comm. 2011).
Remarks. Slightly older than Giganotosaurus andcomparable in size, Tyrannotitan is among the morerecently described carcharodontosaurids (Novas et al.2005). Although the holotype and referred specimen donot include the entirety of the skeleton, the preservedportions of the skull, axial, and appendicular regionsshow a number of carcharodontosaurid features as well asstrong similarities with Giganotosaurus and Mapusaurus.MPEF-PV 1156 also provides evidence of forelimbshortening in derived carcharodontosaurids (Novas et al.2005).
Xuanhanosaurus qilixiaensis Dong, 1984
1984 Xuanhanosaurus qilixiaensis Dong: 213, figs 1–4, pl.1.
Holotype. IVPP V.6729, partial forelimbs and anterior andmiddle dorsal vertebrae.
Diagnosis. Tetanuran with: (1) pronounced posterior ridgeon articular facet of humeral head that overhangs shaft(Rauhut 2003); and (2) dorsal neural spines transverselythick with gently concave lateral surfaces (Rauhut pers.comm. 2007).
Occurrence. Xuanhan County, Sichuan, China; Xiashax-imiao Formation; Middle Jurassic.
Remarks. Known only from an associated forelimband vertebrae, Xuanhanosaurus nevertheless appears topreserve enough morphological information both to distin-guish it from other theropods and to inform its phylogeneticposition. Previously it has been recovered as a basal tetanu-ran (Holtz et al. 2004) and a megalosauroid (Benson 2010a).The robust humerus and wide scapular blade are accompa-nied by a four-digit manus with relatively short metacarpals.This unusual combination of features suggests a basal posi-tion within Tetanurae, and indeed the manus bears someresemblance to those of ‘Szechuanosaurus’ zigongensis andTorvosaurus.
‘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’ hepingen-sis possesses autapomorphies or can only be distinguishedby 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 alter-nately been referred to the genera Yangchuanosaurus (Gao1992, 1999) and Sinraptor (Currie & Zhao 1994; Rauhut2003; Holtz et al. 2004), depending on the perceived impor-tance of particular character states. For example, the propor-tionally long, low skull and very tall dorsal vertebral neuralspines 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 orientedpubic 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 andskeleton (holotype, Y. magnus).
Diagnosis. The original diagnoses of Yangchuanosaurusand its two constituent species (Dong et al. 1978,1983) include features that now characterize ‘sinrap-torids’, allosauroids and tetanurans more generally. Morerecent work (Currie & Zhao 1994) has distinguishedYangchuanosaurus from Sinraptor primarily on the pres-ence of sinraptorid characters together with the absence ofSinraptor autapomorphies (i.e. a unique character combi-nation but no autapomorphies). Yangchuanosaurus has aslightly higher ratio of skull height to length (0.5) thanSinraptor (0.4) and correspondingly has a proportion-ally taller maxilla. The dorsal vertebral neural spines arealso lower (about 1.8 times centrum height comparedto 2.0 in S. dongi) and the centra are relatively longer.Yangchuanosaurus may also exhibit a more pronouncedmargin of the antorbital fossa on the jugal, although thisis difficult to ascertain in some sinraptorid specimens. Weconcur that Yangchuanosaurus can be distinguished fromother sinraptorids, but in the absence of a detailed restudyof the type materials we can offer no new autapomorphiesto define this genus.
Occurrence. Shangyou Reservoir and Hongjiang MachineFactory, near Yongchuan, Yongchuan County, Sichuan,China; Shangshaximiao Formation; Oxfordian–earlyKimmeridgian, Late Jurassic.
Remarks. The original and emended diagnoses ofYangchuanosaurus shangyouensis and Y. magnus differ-entiated these species primarily on the basis of size (Donget al. 1978, 1983). In addition, Dong et al. (1983) notedthat the maxilla of Y. magnus housed an additional fenes-tra within the antorbital fossa, whereas Y. shangyouensispossessed only a fossa in this location. Given the tendenciestowards increased pneumatization in larger (older) individ-uals among theropods (Wedel 2003; although see Rauhut &Fechner 2005; Brusatte et al. 2009b) and the variability ofpneumatic features even within a single individual (Zhao &
Currie 1994), we consider this likely to be an intraspecific,possibly ontogenetic, variation. The apparent difference incervical vertebral morphology (ventral keels absent in Y.shangyouensis, present in Y. magnus) may be a functionof comparing different positions within the column. There-fore, the holotypes of the two species of Yangchuanosaurusare effectively identical, and indeed they would have thesame patterns of character codings in our matrix. Giventheir provenance and the high degree of similarity betweenthese specimens, we consider them to represent a singlespecies.
Phylogenetic analysis
CharactersThis study employed a total of 351 characters derived froma combination of previous phylogenetic studies (see onlineSupplementary Table 1) and direct study of specimensby the authors. Numerous previously described charac-ters were not employed in this study, and it would be agargantuan task to discuss these individually. In general,‘excluded’ characters were: (1) parsimony uninformativefor our taxon set; (2) redundant or subsumed into exist-ing characters; or (3) not explicable or observable by thepresent authors. All included characters were scored basedon direct examination of fossil materials or casts, except inthose cases where key specimens no longer exist or wherewe could not visit for study. In total, more than 90% ofincluded taxa were examined directly by one or more ofthe authors. Character descriptions are listed in Appendix 3(see Supplementary Material online). The taxon-charactermatrix is given in Appendix 2, and executable version isavailable online as Supplementary File 3.
Phylogenetic methodsThe 351 characters used in this analysis were unorderedand equally weighted, and scored using Mesquite 2.72(Maddison & Maddison 2009). The resulting matrix wasimported into TNT 1.1 (Goloboff et al. 2008). Given thelarge size of this matrix, we opted for a heuristic searchusing the ‘New Technology’ options. These included thedefault settings for sectorial, ratchet, tree drift and treefusion, using a driven search that stabilized consensus twicewith a factor of 25. Subsequently, we subjected the result-ing most parsimonious trees (MPTs) to tree bisection andreconnection (TBR) branch swapping. The trees resultingfrom both search iterations were examined using strict andAdams consensus (‘combinable components’ in TNT). Weconfirmed the results of this analysis using PAUP∗ 4.0b10(Swofford 2002), employing the parsimony ratchet (Nixon1999) implemented in PAUPRat (Sikes & Lewis 2001) andTBR branch swapping on the resulting trees.
We isolated ‘wildcard’ taxa using the Adams consensustree. ‘Wildcards’ are taxa that take multiple phylogenetic
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placements with equal parsimony, thus resulting in a poorlyresolved strict consensus. Pruning wildcards from the setof MPTs and then calculating the strict consensus resultsin a ‘strict reduced consensus’. This strategy summarizesrelationships among non-wildcard taxa that are otherwiseconcealed by inclusion of wildcards (Wilkinson 2003). Weperformed successive iterations of our search strategy onthe same character matrix after having removed these taxa.
Following the initial analyses, we determined branchsupport using the ‘Decay Index PAUP File’ function ofMacClade (Maddison & Maddison 1992) combined withthe parsimony ratchet implemented using PAUPRat inPAUP∗ to determine the shortest tree length lacking eachclade that was originally recovered by our analysis.
Stratigraphic fit methodsWe employed three commonly used methods to assess thestratigraphic fit of our phylogenetic results. We used aSpearman rank correlation to test the relationship betweenage rank (using stages, numbered from oldest to youngest)and clade rank (numbering nodes upwards from the base ofthe tree), although we did not collapse the cladogram intoa ‘comb’ topology. The Stratigraphic Consistency Index(SCI; Huelsenbeck 1994) was used to assess the number ofnodes that were consistent with the stratigraphic placementof the oldest included taxa. Finally the Relative Complete-ness Index (RCI; Hitchin & Benton 1997) measures theamount of record missing based on the current phylogeny.
Results
Tree length and topologyThe initial TNT analysis produced 91 most parsimonioustrees, each of 1020 steps; subsequent TBR branch swappingresulted 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 rescaledconsistency index (RC) = 0.2932. Excluding parsimonyuninformative characters, the CI was 0.2404 and the HIwas 0.5796. Results were identical when PAUP∗ was usedto perform the search, but more than 400,000 MPTs wererecovered. A strict consensus of these trees produced thecladogram in Fig. 7A.
This phylogeny provides significant resolution among themajor tetanuran clades that confirms the successive place-ment of coelophysoids, ceratosaurs and tetanurans withinTheropoda. Dilophosaurus is recovered as a coelophysoid,in contrast to a recent study (Nesbitt et al. 2009), whereasCryolophosaurus and ‘D.’ sinensis are more derived. Teta-nurae is monophyletic and includes most of its traditionalsubclades.
Within Tetanurae we recovered: (1) a basal clade, Mega-losauroidea (= Spinosauroidea, see below), which includes
spinosaurids, megalosaurids and a third, more basal clade;(2) a more derived Allosauroidea, including allosaurids,sinraptorids (= metriacanthosaurids, see below), neovena-torids and carcharodontosaurids; and (3) Coelurosauria,here represented by Compsognathus, Ornitholestes andProceratosaurus, the sister taxon to Allosauroidea.
Megalosauroidea is composed of three main clades.Among them, Spinosauridae includes Spinosaurinae(Spinosaurus, Irritator and Angaturama) and Baryonychi-nae (Baryonyx and Suchomimus). Its sister lineage Mega-losauridae is a diverse assemblage that includes at least twoclades, Megalosaurus + Torvosaurus + Duriavenator anda poorly resolved second group (Dubreuillosaurus, Afrove-nator, Piveteausaurus, Leshansaurus, Magnosaurus andPoekilopleuron); Eustreptospondylus and Streptospondylusare unresolved within Megalosauridae + Spinosauridae. APiatnitzkysaurus + Marshosaurus + Condorraptor cladeis found to be basal within Megalosauroidea.
Among the allosauroids, ‘sinraptorids’ are the most basalgroup. The clade is more diverse than has previously beenassessed, with six taxa aggregated into two groups (Metri-acanthosaurus + Sinraptor dongi + Siamotyrannus +‘Y.’ hepingensis and Yangchuanosaurus shangyouensis +‘S.’ zigongensis + CV 00214) plus two unresolved basalforms (Shidaisaurus and Xuanhanosaurus). Allosauridaeexists only as Allosaurus + Saurophaganax, the sisterlineage to Carcharodontosauria (Carcharodontosauridae +Neovenatoridae). Several recently discovered forms makethis latter the most diverse allosauroid clade. Neovena-toridae comprises Neovenator, Chilantaisaurus, Aerosteon+ Megaraptor, and Australovenator + Fukuiraptor. Theremaining taxa (Eocarcharia, Concavenator, Shaochilong,Acrocanthosaurus, Mapusaurus, Tyrannotitan, Carchar-odontosaurus and Giganotosaurus) form Carcharodon-tosauridae.
This analysis recovered several individual taxa in well-resolved positions outside major clades or ingroups. Amongthem, Cryolophosaurus and ‘D.’ sinensis are basallyplaced ‘stem’ tetanurans; Monolophosaurus and Chuan-dongocoelurus occupy similar but slightly more derivedpositions. Interestingly, Lourinhanosaurus is allied withCoelurosauria.
An Adams consensus (Fig. 7B) identified Poekilopleu-ron, Streptospondylus and Xuanhanosaurus as ‘wildcard’taxa. Deletion of Streptospondylus resulted in a four-fold reduction to 105,750 unique trees and the resolu-tion of Eustreptospondylus as the most basal megalosaurid.Subsequent deletion of Xuanhanosaurus resulted in anadditional five-fold reduction to 21,150 unique trees andresolution of Shidaisaurus as a basal representative ofMetriacanthosaurinae; eliminating Poekilopleuron resultedin 4050 trees, with no further improvement in resolution(Fig. 8A).
Finally, we conducted an Adams consensus on the 4050trees 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 significantlyredefined clades resulting from this analysis. Numbers to the left of the nodes indicate unambiguous character support, those to the rightshow 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 mostparsimonious 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 actingas ‘wildcards’, but their removal does not improve resolu-tion beyond that achieved by deleting one taxon from anypolytomy.
Branch support and taxon instabilityCharacter support for each node is reported in onlineSupplementary File 3. Branch support is low (≤2) for manynodes, but several clades (Coelophysidae, Neotheropoda,Averostra, Ceratosauria, Spinosauridae) show significantsupport (> 5). However, in many instances the low supportis due to the uncertain position of a single taxon. Theseare typically fragmentary or otherwise poorly known formswhose instability gives a spuriously low confidence valuefor the overall scheme of relationships.
For example, a single added step allows the follow-ing placements: Chilantaisaurus as a basal carcharo-dontosaurid close to Concavenator; Shidaisaurus as abasal metriacanthosaurid or the sister taxon to Metria-canthosaurus; Monolophosaurus as a basal megalosauroid;and Cryolophosaurus outside Averostra or just above ‘D.’sinensis. Lourinhanosaurus can be recovered as a basalallosauroid or a basal avetheropod with the addition of twosteps.
In addition, we recover Poekilopleuron as an afrovena-torine megalosaurid, but with a single additional step itcan be recovered as a piatnitzkysaurid, elsewhere withinMegalosauridae, or at several possible positions withinAllosauria. This effectively reduces support for the entireavetheropod ‘stem’, many allosaurian ingroups, and muchof Megalosauroidea down to 1. A significant improve-ment in overall support is actually achieved if Poekilopleu-ron is removed from the analysis and branch support isrecalculated. In this case, support for several nodes risessignificantly, including Allosauria and Allosauroidea (to3), and Metriacanthosauridae and Carcharodontosauria (to4), while most other nodes now have a support of at least 2.Given how fragmentary the only known specimen of Poek-ilopleuron is, we consider its placement within Afrovena-torinae to be extremely tentative, and realistically it couldrepresent another megalosaurian or even an allosaurian.
Taxonomic and nomenclatural implicationsThe topologies produced by our analyses indicate severalchanges to existing taxonomic usage. These are describedbelow, and summarized in Appendix 1.
The family Megalosauridae was established in Fitzinger(1843; non Huxley 1870) in accordance with ICZN Article11.7.2. It takes priority over Spinosauridae Stromer, 1915and Torvosauridae Galton & Jensen, 1979; therefore Mega-losauroidea is a senior synonym of Spinosauroidea andTorvosauroidea. It is used here to include all taxa closerto Megalosaurus than to Allosaurus or Tyrannosaurus.Within this clade, Spinosauridae and Megalosauridae are
most closely related; we append the existing name Mega-losauria Bonaparte, 1850 to this node. Megalosaurinae canbe defined as all megalosaurids closer to Megalosaurus thanto Afrovenator.
Metriacanthosaurus is the type genus of Metriacan-thosauridae Paul, 1988a, which was defined only briefly butwith purportedly differential characters (in accordance withICZN Article 13.1.1). The robust recovery of Metriacan-thosaurus within Sinraptoridae Currie & Zhao, 1994 placesthe latter family name into synonymy with the former.
Two lineages are evident within Metriacanthosauri-dae, but individual taxa are not interrelated in amanner that supports all previous species assignments.For example, we recover a sister-taxon relationshipbetween Yangchuanosaurus shangyouensis and specimenCV 00214, with the type of ‘Szechuanosaurus’ zigongen-sis as outgroup to this pair. This contradicts the sugges-tion that the latter two specimens pertain to a singlespecies (Chure 2001a). Rather, we assign CV 00214 toY. shangyouensis given its close similarity and deriva-tion from the same stratum and geographic region. Thuswe agree with Paul (1988a) that it is congeneric withYangchuanosaurus shangyouensis but do not agree thatthe differences warrant specific separation. We acknowl-edge the close relationship between ‘Szechuanosaurus’zigongensis and Yangchuanosaurus by referring the formerto Yangchuanosaurus as Y. zigongensis but retain it as adistinct species given its earlier provenance. The nameYangchuanosaurus can be applied to this entire clade.
However, ‘Y.’ hepingensis is more closely related toSinraptor than to Yangchuanosaurus, supporting its recentreferral to the former genus (Currie & Zhao 1994). S. hepin-gensis and S. dongi belong to a lineage that also includesMetriacanthosaurus and Siamotyrannus, and can be definedas follows.
Tetanurae Gauthier, 1986Allosauroidea Currie & Zhao, 1994Metriacanthosauridae Paul, 1988a
Metriacanthosaurinae (Paul 1988a) subfam. nov.
Type genus. Metriacanthosaurus Walker, 1964.
Included taxa. Metriacanthosaurus parkeri (Huene,1923) Walker, 1964; Siamotyrannus isanensis Buffetautet 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 followingsynapomorphies: (1) anteroventral border of maxillaryantorbital fossa demarcated by raised ridge (22(1); also inEoraptor, Coelophysidae, Masiakasaurus, Marshosaurusand Compsognathus); (2) pronounced ventral keel on
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anterior dorsal vertebrae (184(1)); also in Condorrap-tor, Piatnitzkysaurus, Carcharodontosaurus and manymegalosaurians); (3) straight posterior margin of iliacpostacetabular 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 Coelophysi-dae, Eustreptospondylus, Afrovenator, Megalosaurus andCompsognathus); and (5) bulbous fibular crest on tibia(323(1)); also in ‘Dilophosaurus’ sinensis).
Definition. All metriacanthosaurids more closely relatedto Metriacanthosaurus than to Yangchuanosaurus.
Allosauridae is reduced to Allosaurus + Saurophaganaxbut holds a stable place as the sister taxon to Carcharodon-tosauria. We suggest using the existing name AllosauriaPaul, 1988a for this node, in parallel with the nomenclatu-ral arrangement outlined above for megalosauroids.
The position of ‘D.’ sinensis as distinct from D. wetherilliis likely secure, as the two taxa are well separated in ourresults, and therefore this species requires assignment toa 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 Mega-losauroidea. These taxa show numerous morphologicalsimilarities as well as distinctions from other mega-losauroids, and we consider it useful to provide a namefor this grouping.
Tetanurae Gauthier, 1986Megalosauroidea (Fitzinger, 1843)
Piatnitzkysauridae fam. nov.
Type genus. Piatnitzkysaurus Bonaparte, 1979.
Included taxa. Piatnitzkysaurus floresi Bonaparte, 1979;Marshosaurus bicentesimus Madsen, 1976b; Condorraptorcurrumili Rauhut, 2005a.
Diagnosis. Megalosauroids possessing the followingsynapomorphies: (1) two parallel rows of nutrient foraminaon lateral surface of maxilla (21(1)); also in Eocarchariaand Shaochilong); (2) vertically striated or ridged paraden-tal plates (140(1)); also in abelisaurids, Megalosaurus andProceratosaurus); (3) reduced axial parapophyses (166(1));also in coelophysoids, Eustreptospondylus and Afrovena-tor); (4) anteriorly inclined posterior dorsal neural spines(192(1)); in parallel with Allosauroidea); and (5) canteddistal humeral condyles (236(1)); also in Poekilopleuron,Allosauridae and Fukuiraptor). In addition, among tetanu-rans these taxa show reversals to the primitive conditionfor the following features: (1) a short or absent anteriormaxillary ramus (12(0)); (2) moderate development of axialdiapophyses (167(0)); and (3) no axial pleurocoels (168(0)).
Definition. All megalosauroids more closely related toPiatnitzkysaurus than to either Spinosaurus or Mega-losaurus.
Finally, we have recovered a novel clade withinMegalosauridae that includes six taxa: Piveteausaurus,Leshansaurus, Afrovenator, Dubreuillosaurus, Poekilo-pleuron and Magnosaurus. No existing name can be appliedto this clade, so we supply the following.
Tetanurae Gauthier, 1986Megalosauroidea Fitzinger, 1843Megalosauridae Fitzinger, 1843Afrovenatorinae subfam. nov.
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 tenta-tively Poekilopleuron bucklandii Eudes-Deslongchamps,1837.
Diagnosis. Megalosauroids possessing the followingsynapomorphies: (1) squared anterior margin of maxillaryantorbital fenestra (23(1)); also in coelophysoids, Irrita-tor, Concavenator and Eocarcharia); and (2) puboischiadicplate broadly open along midline (281(2)); in parallel withAvetheropoda).
Definition. All megalosaurids more closely related toAfrovenator than to Megalosaurus.
Finally, we establish the new name Orionides for the nodecomprising Megalosauroidea, Avetheropoda, their mostrecent common ancestor, and all its descendants. The namealludes to Orion, the giant hunter of Greek mythology (andto the alternate name for the constellation Orion, Alektropo-dion, or ‘chicken foot’).
Fragmentary taxaAs with our analysis of ceratosaurs (Carrano & Sampson2008), we did not include every potential basal tetanuranin this study; several were excluded because they were toofragmentary or provided no unique character combinations.The phylogenetic determinations discussed below are madeto the most exclusive level possible. Although many of theseforms cannot be identified beyond a rather general level (i.e.Allosauroidea), they nonetheless add important temporaland geographic data to the story of tetanuran evolution(Fig. 9). In keeping with the scope of this study, we haveexcluded coelurosaurs from the discussion below, and donot discuss every report of a theropod tooth unless it hasbeen previously assigned to a specific basal tetanuran taxon.
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Jurassic CretaceousEarly Middle Late Early Late
He Si Pl To Aa Bj Bt Cl Ox Ki Ti Be Va Ha Ba Ap Al Ce Tu Co Sa Ca Ma
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EUR
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AcrocanthosaurusSaurophaganax
Allosaurus, Torvosaurus
Dubreuillosaurus, Poekilopleuron
Magnosaurus
Eustreptospondylus
Piveteausaurus, Streptospondylus
Metriacanthosaurus
Lower Lias
Condorraptor, Piatnitzkysaurus
Afrovenator
‘D.’ sinensis
Cryolophosaurus
Baryonyx
Cristatusaurus, Suchomimus
Spinosaurus
Carcharodontosaurus
Tyrannotitan
Angaturama, Irritator
Neovenator, Baryonychinae
Leshansaurus, S. hepingensisY. shangyouensis
GiganotosaurusMapusaurus
Gasosaurus, Kaijiangosaurus,Shidaisaurus, Xuanhanosaurus,
Y. zigongensis
AllosaurusTorvosaurus
Megaraptor
‘A.’ tendagurensis, Carcharodontosauria
‘Saltriosaurus’
Jobu
Pinilla de los Moros, Blesa
S. girardi
Artoles
Unquillosaurus
OrkoraptorAlcântaraCerro Lisando
‘M.’ inexpectatus
Adamantina
Allen
Cabao
Wadi MilkMugher Mudstone
Siamosaurus, Siamotyrannus
Hidden Lake
Fukuiraptor
‘A.’ sibiricus, Embasaurus
BecklespinaxValdoraptor
MorellaHastings
Villar del Arzobispo
S.dongi
Chuandongocoelurus, MonolophosaurusBalabansai
Montmirat
Aerosteon
Australovenator
Chilantaisaurus, Shaochilong
Eocarcharia
Cruxicheiros, Iliosuchus, Megalosaurus
Marshosaurus
Erectopus
Khok Kruat
‘S.’ cuvieriJet Rock
‘M.’ ingens
‘M.’ pannoniensis
Eumeralla
Phu Kradung
Arundel
Enciso
Kimmeridge Clay
Wessex
Kirkwood
Cloverly
Marília
Duriavenator
Rapator
Kelmayisaurus
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 thedistal 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 istoo fragmentary to be assigned to a known taxon or identi-fied as a distinct form, especially considering that this boneis not particularly diagnostic among non-coelurosauriantheropod clades. Its morphology is quite similar to thatof Allosaurus and Neovenator but also to more primitiveforms such as Afrovenator and Torvosaurus. Holtz et al.(2004) referred to it as Chilantaisaurus sibiricus, but it isbest identified as Theropoda indet.
Allosaurus? tendagurensis Janensch, 1925. A single tibiafrom 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 elon-gate, the proximal and distal ends are sufficiently wellpreserved. As suggested by Rauhut (2005b), this speci-men clearly pertains to a tetanuran; it lacks the distinc-tive ceratosaur cnemial crest, the distal end bears a markedbuttress for the (presumably laminar) astragalar ascend-ing process, and the distal profile is transversely elongate.However, the specimen is not particularly similar to that ofAllosaurus among tetanurans and cannot be more specif-ically identified. The assignment of isolated caudal verte-brae from Quarry TL to this taxon (Janensch 1925) is notjustified.
Becklespinax altispinax (Paul, 1988a) Olshevsky, 1991.Three dorsal vertebrae (NHMUK R1828; Fig. 10) fromthe Hastings Beds Group (late Berriasian–Valanginian) ofBattle, England constitute the controversial type specimen
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The phylogeny of Tetanurae 253
Figure 10. Posterior dorsal vertebrae (NHMUK R1828) of Beck-lespinax altispinax (Paul, 1988a) in left lateral view. Abbrevia-tions: hs, hyposphene; pb, pathological bone; poz, postzygapoph-ysis; prz, prezygapophysis. Scale bar = 10 cm.
of this taxon. Originally, these materials were condition-ally assigned to Megalosaurus dunkeri Dames, 1884 underthe new genus Altispinax Huene, 1923. Specifically, Huene(1926a, p. 78) remarked that if these vertebrae belonged tothe same animal as the holotype tooth of M. dunkeri, thenthe resulting taxon could be called Altispinax dunkeri. Asthis tooth is non-diagnostic within Theropoda (see below),and in any case cannot be assigned to the same taxon asthe vertebrae, NHMUK R1828 was made the holotype of anew taxon (Paul 1988a; Olshevsky 1991).
The vertebrae are unusual in possessing highly elon-gate neural spines, which has led many workers (Walker1964; Romer 1966; Carroll 1988; Kurzanov 1989) to assignBecklespinax to Spinosauridae. However, they show impor-tant differences with the vertebrae of both spinosaurids andthe long-spined carcharodontosaurids. Unlike spinosaurids,the arches lack accessory laminae either below or abovethe transverse process. Unlike carcharodontosaurids (andTorvosaurus), the centra are poorly or non-pneumatized,
showing only weak lateral indentations that are widelypresent among amniotes, and the lateral surfaces of theneural spines do not bear deep longitudinal troughs (derivedcarcharodontosaurids only). The spines are unusually thick,approximately as broad mediolaterally as they are longanteroposteriorly, and widen considerably toward theirapices. They also show evidence of a healed break as wellas some ossification of the interspinous (presumably liga-mentous) soft tissues. The centrum proportions are similarto those of many non-coelurosaurian tetanurans. Other thanto exclude Becklespinax from Spinosauridae and Carchar-odontosauridae, we cannot place it more specifically withinTetanurae. Its exclusion from these two clades implies theindependent derivation of elongate dorsal neural spines atleast three times within Theropoda.
Cristatusaurus lapparenti Taquet & Russell, 1998.Cristatusaurus lapparenti was the first spinosauriddescribed from the same formation (Elrhaz Formation;Aptian–Albian) and general location as Suchomimus(Taquet 1976; Taquet & Russell 1998). It has been widelyconsidered a nomen dubium (Sereno et al. 1998; Sues etal. 2002; but see Allain 2002) because the holotype mate-rial (MNHN GDF 366) is extremely fragmentary and theaccompanying diagnosis very general, a conclusion withwhich we agree. Cristatusaurus can only be identified as anindeterminate baryonychine spinosaurid. However, we alsoconsider it unlikely that two baryonychines are present atGadoufaoua, and therefore Cristatusaurus and Suchomimusalmost certainly represent the same animal.
Cruxicheiros newmanorum Benson & Radley, 2010. Arecently described fragmentary skeleton from the LowerBathonian of Cross Hands Quarry, Warwickshire repre-sents a large, early basal tetanuran that is distinct fromMegalosaurus and SDM 44.19 (see below). The specimens(WARMS G15770, G15771) preserve at least one autapo-morphy as well as numerous features differentiating Cruxi-cheiros from coeval forms, but it is sufficiently incompletethat its placement within Tetanurae is uncertain (Benson &Radley 2010).
Embasaurus minax Riabinin, 1931. Based only on a pairof dorsal vertebral centra from the Neocomian (LowerCretaceous) of Kazakhstan, Embasaurus minax was orig-inally assigned to Megalosauridae (Riabinin 1931). LikeBecklespinax, the absence of pneumatic foramina (pleu-rocoels) in the posterior dorsal centra seems to excludethis taxon from Carcharodontosauria. The centra areamphiplatyan but the neural arches are not preserved.Embasaurus is here considered a nomen dubium, and thereis little additional information to identify it more specifi-cally than as Theropoda indet.
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Erectopus superbus (Sauvage, 1882) Huene, 1923. Thepartial skeleton forming the holotype of Megalosaurussuperbus was collected from the lower Albian (Douville-iceras mammilatum Zone) phosphatic ‘La Penthieve Beds’of Bois de la Penthiere, near Louppy-le-Chateau, Meuse,France (Sauvage 1882; Huene 1923; Allain 2005b). Thematerial pertains to a medium-sized theropod and consistsof a partial maxilla, incomplete manus, left femur andtibia, 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 limbelements. The anteromedial (palatal) process of the maxillais located ventrally, adjacent to the dorsal margins ofthe paradental plates, as in Marshosaurus and Sinrap-tor. The femur suggests avetheropod affinities as it lacksa proximal articular groove (Hutchinson 2001; Benson2010a), and the condyles are separated by a shallowanteroposteriorly oriented trough on the distal surface.The distinct, step-like supraastragalar buttress of the tibiais distinct from the reduced buttress of carcharodon-tosaurians. We suggest that Erectopus represents a non-carcharodontosaurian allosauroid, possibly a metriacan-thosaurid.
Gasosaurus constructus Dong, 1985. Gasosaurus isbased on vertebrae, a humerus, and highly abraded andreconstructed pelvic and hind limb material from theMiddle Jurassic Xiashaximiao Formation of Dashanpu,Sichuan, China and was originally described as a mega-losaurid. Holtz (2000) recovered Gasosaurus as a basalcoelurosaur but Currie (in Holtz et al. 2004) stated thatnew specimens indicated it was a primitive ‘carnosaur’(allosauroid). Many potentially informative features of theholotype skeleton (IVPP V7265) are difficult to assessbased on published descriptions and images. The taxonrepresents a tetanuran based on the presence of a pubicpeduncle of the ilium that is substantially larger than theischial peduncle. Examination of casts reveals that the lessertrochanter does not rise above the level of the femoral head(contra Holtz 2000; Holtz et al. 2004). Instead, the prox-imal portion of the femoral head is broken and the lessertrochanter reaches approximately midlevel of the head as innon-coelurosaurian tetanurans. For now Gasosaurus is bestregarded as having an uncertain position within Tetanu-rae and probably outside Coelurosauria; however, detailedrestudy of the holotype is underway (D. Hone pers. comm.).
Iliosuchus incognitus Huene, 1932. Iliosuchus includes asmall partial ilium from Stonesfield, the topotype localityof Megalosaurus bucklandii (NHMUK R83; Taynton Lime-stone Formation, Bathonian, Middle Jurassic, England) andtwo referred ilia (OUMNH J.29780 [Galton 1976] andOUMNH J.29871 [Foster & Chure 2000]). The presenceof a swollen ridge on the lateral surface of the ilium hasbeen considered the diagnostic feature of this taxon. It hasbeen used to unite the genus with the basal tyrannosauroid
Stokesosaurus clevelandi from the Late Jurassic of NorthAmerica (Galton 1976), but this feature has a wider distri-bution among theropods. Benson (2009b) observed thatit was also present in M. bucklandii, suggesting that I.incognitus showed no diagnostic features and should beconsidered a nomen dubium, representing either an inde-terminate avetheropod or a juvenile specimen of M. buck-landii. OUMNH J.29871 is distinct from the holotype ofI. incognitus because it has a vertical, not posterodorsallyinclined, median ridge, larger size, and several small ‘acces-sory’ ridges anteriorly and posteriorly on the lateral surface.However, it cannot be identified other than as Tetanuraeindet.
Kaijiangosaurus lini He, 1984. Seven cervical vertebraeconstitute the holotype of Kaijiangosaurus lini (CCG20020), which was discovered in the Middle JurassicXiashaximiao Formation of Sichuan Province, China (He1984). These elements are not elongate and have a flatanterior surface and a concave posterior surface, which arenot significantly offset from one another. The neural spineis anteroposteriorly short and slightly posteriorly inclined.These features suggest placement as a basal tetanuran orbasal averostran. Referred specimens include a tooth, jugal,two dorsal and seven caudal vertebrae, scapula and cora-coid, humerus, proximal ulna, partial manus, femur, partialtibia and fibula, tarsus, and incomplete pes. Unfortunatelyit is not clear whether they collectively represent a singletaxon. Restudy of these materials is needed.
Kelmayisaurus petrolicus Dong, 1973. A left dentary andmaxilla constitute the holotype (IVPP V 4022) of this taxon,which was recently redescribed, diagnosed, and assigned toCarcharodontosauridae (Brusatte et al. 2011). The presenceof Kelmayisaurus in the Lianmugin Formation (Early Creta-ceous) of China emphasizes the lengthy tenure of carchar-odontosaurids in Asia.
Orkoraptor burkei Novas, 2008. Orkoraptor is based onfragmentary remains from the Pari Aike Formation (?Maas-trichtian) near Los Hornos Hill, Santa Cruz Province,Argentina. These were originally identified as a coelurosaurbut have now been demonstrated to pertain to a megarap-toran allosauroid (Benson et al. 2010) based on the presenceof proximal caudal pleurocoels, neural arch foramina andlaminae and an Aerosteon-like postorbital. The affinities ofOrkoraptor within Megaraptora remain unresolved due tothe incomplete nature of the holotype and only specimen(MPM-Pv 3457).
Poekilopleuron schmidti Kiprianow, 1883. This taxon isbased on a three partial ribs and a distal humerus fromthe Albian–Cenomanian Sekmenevsk Formation of Tuskar,near Meshkovo, in western Russia. Interestingly, Kipri-anow (1883) presented several histological sections of thesebones showing details of the internal microstructure, one
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The phylogeny of Tetanurae 255
of the earliest examples for dinosaurs. The bones them-selves are quite fragmentary and largely uninformative. Thehumerus seems to exhibit a hollow interior (also present insmall ornithopods, but these are smaller than P. schmidti)and the distal end is flattened anteroposteriorly. The radialcondyle and ectepicondyle are broken but the ulnar condyleis intact and wraps slightly onto the posterior surface. Thereis a small entepicondyle on the medial side. At present wecan only tentatively confirm the theropod nature of thiselement, and the species cannot be assigned to the genusPoekilopleuron.
Rapator ornitholestoides Huene, 1932. The holotype ofRapator comprises only a left metacarpal I (NHMUKR3718), from the Griman Creek Formation (Albian) ofLightning Ridge, Australia. It was originally consideredto pertain to a coelurosaur similar to Ornitholestes (Huene1932), and more recently as manual phalanx I-1 from analvarezsaurid (Holtz et al. 2004). However, Agnolin et al.(2010) noted similarities with metacarpal I of Australove-nator and Megaraptor, and suggested that Rapator wasa nomen dubium belonging to Megaraptora. We agreewith this morphological assessment, but in the absence ofcomparison to the same element in Neovenator (currentlyunknown), we cannot place it more specifically than Neove-natoridae indet.
Siamosaurus sutheethorni Buffetaut & Ingavat, 1986.Siamosaurus is based on isolated teeth (TF 2043a-i)from the Sao Khua Formation (Barremian–Aptian) of PhuWiang, Thailand that resemble those of spinosaurids inexhibiting fluted enamel, a less recurved profile and rela-tively rounded cross section (although much less so thanin some spinosaurids). The teeth do resemble those ofspinosaurids but not enough material is known to supportretention of a specific taxon, and their theropod affinitieshave 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 KhokKruat Formation of Thailand may eventually permit a moresecure identification (Buffetaut et al. 2004, 2005; Milneret 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 ofChipping Norton, Oxfordshire, England. These were asso-ciated with a “broad flat bone” and fragments of longbones (Owen 1842, p. 90). It was subsequently trans-ferred to Megalosaurus by Huene (1908, 1932) as the newcombination M. cuvieri. Notably, the holotype specimensof Streptospondylus altdorfensis, Eustreptospondylus andPiveteausaurus were all at one time referred to S. cuvieri(Nopcsa 1906; Piveteau 1923; Huene 1932). Unfortunately,
the holotype and referred specimens of S. cuvieri were neverfigured and are now presumed lost.
The holotype vertebra is likely to have been an anteriordorsal because “the articular surfaces of the ribs are, asusual, 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 openingthat deeply invaginated into the body of the bone. Typicallaminae were present on the base of the neural arch. Theneural spine exhibited pronounced attachments for the inter-spinous ligaments along its anterior and posterior edges.The tooth was described as conical and hollow, and maynot pertain to a theropod. The remaining specimens indi-cate an indeterminate tetanuran theropod that cannot bereferred to the genus Streptospondylus.
Owen also mentioned a second specimen, a posteriordorsal vertebra from the “jet-rock (lias shales)” (Jet RockFormation, Lower Toarcian; Howarth 1980) of Whitby inthe collection of Mr Ripley, as “referable to the presentgenus” (Owen 1842, p. 90). It too is lost but was apparentlysimilar in basic morphology to the type specimens of ‘S.’cuvieri, although more complete. It was clearly opistho-coelous and preserved more of the neural arch includingthe zygapophyses (“oblique processes”; Owen 1842, p. 90).The presence of a large, single pneumatic foramen in thecentrum is a synapomorphy of Tetanurae and the Toarcianspecimens may be one of the earliest records of the clade,but as no diagnostic features were described they remainTetanurae indet.
Suchosaurus cultridens (Owen, 1841) Owen, 1842. Thistaxon was originally named Crocodilus cultridens for asingle tooth (NHM R36536) that Owen (1841 in Owen1840–1845) thought belonged to a crocodilian; he laterremoved it to the new genus Suchosaurus (Owen 1842).Only much later, following the discovery of Baryonyx, wasthe material re-examined and referred to the Baryonychinae(Buffetaut 2007, 2010; Mateus et al. 2011). The tooth itselfis not diagnostic beyond the family level, and we considerit to be a nomen dubium.
Suchosaurus girardi Sauvage, 1897–1898. The materialon which Sauvage based Suchosaurus girardi is slightlymore complete than that used by Owen to describe the typespecies, S. cultridens, and includes a jaw fragment withteeth. These specimens bear several features in commonwith baryonychines (Buffetaut 2007, 2010; Mateus et al.2011) but no diagnostic qualities of their own. S. girardi istherefore a nomen dubium; it cannot be formally referredto the later-named genus Baryonyx without subsuming thatname into Suchosaurus. However, it derives from the LowerBarremian Papo Seco Formation of Portugal, where Bary-onyx is known from other material (Mateus et al. 2011) andmay 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 JurassicKuangyuan Formation of Sichuan Province, China, butthese are not diagnostic below the level of Theropoda. Asthis is the type species of the genus Szechuanosaurus, thattaxon cannot be diagnosed and should not be used to receiveadditional species.
Unquillosaurus ceibalii Powell, 1979. Based on anisolated pubis from the Los Blanquitos Formation (lateSantonian–?Campanian) of Arroyo El Morterito (Campa-nian), Salta, Argentina, Unquillosaurus has been difficultto place phylogenetically. Originally described as the leftpubis of an indeterminate large theropod (Powell 1979), itwas recently redescribed as belonging to a giant manirap-toran (Novas & Agnolin 2004).
In contrast to all previous workers, we are certain thatthis specimen (PVL 3670-11) is a right pubis. The ‘lateralcrest’ (“cresta lateral”; Powell 1979, fig. 7) is the pubicapron, which bears a rugose contact surface for the oppos-ing element at approximately midshaft. The obturator fora-men, which is genuinely open, is flush with the lateralsurface but inset from the medial surface at the proximalend. 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 lateralmargin is nearly straight but the medial margin is sinuous,suggesting the presence of a foramen between the pubeslocated proximal to the contacting distal ends. The distalend itself is eroded and broken, and lacks an anterior expan-sion although this may have been present when complete. Itmay also have been expanded considerably posteriorly; theposterior margin of the shaft forms an acute angle beforeterminating at a break.
In all, the morphology of this bone resembles that ofGiganotosaurus in many respects, and we suggest that itbelongs to a carcharodontosaurid. It is possibly distinctenough 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 iden-tified as a specimen of Hylaeosaurus (Owen 1858) andlater named as a species of Megalosaurus (Lydekker 1889).Its age (Valanginian) alone makes it an unlikely memberof this genus, and Olshevsky (1991) removed it to a newform, Valdoraptor. Although there may be other distinc-tive features about the holotype (Naish & Martill 2007), itis probably an avetheropod based on the trapezoidal crosssection of metatarsal III, a unique, unambiguous synapo-morphy in our analysis.
Wakinosaurus satoi Okazaki, 1992. This taxon is basedon a single tooth (KMNH VP 000,0016) from the Sengoku
Formation (Hauterivian–Barremian) of Fukuoka, Japan,that was distinguished by the presence of longitudinal stria-tions. Unlike other teeth with similar features, it is stronglylaterally compressed. Okazaki (1992, p. 88) referred Waki-nosaurus to Megalosauridae and suggested that it mightshare affinities with the tooth taxon Prodeinodon kwang-shiensis. Although the tooth is distinctive, it cannot beassigned either to Megalosauridae or to any particular cladewithin Theropoda at this time.
Species previously assigned to the genusMegalosaurusA host of species ranging in age from Late Triassicthrough Late Cretaceous has been placed within the genusMegalosaurus, but presently we consider only the typespecies M. bucklandii to be properly assigned to it (Benson2009a, 2010a). All other taxa originally referred to Mega-losaurus that are represented by diagnostic materials havesince been assigned to other genera. They have beenincluded in our analysis or discussed in the previous sectionand are only listed here: Carcharodontosaurus saharicus,Dilophosaurus wetherilli, Duriavenator hesperis, Erec-topus superbus, Magnosaurus nethercombensis, Majun-gasaurus crenatissimus, Proceratosaurus bradleyi andValdoraptor oweni. Betasuchus bredai is considered tobelong to Ceratosauria (Carrano & Sampson 2008).
A number of theropod taxa have been included withinMegalosaurus in previous taxonomic revisions althoughthey were not originally named as species of thatgenus. These include Antrodemus valens (Nopcsa 1901),Ceratosaurus nasicornis (Cope 1892), Deinodon horridus(Nopcsa 1901), Dryptosaurus aquilunguis (as a subgenusof Megalosaurus; Deperet & Savornin 1928), Iliosuchusincognitus (Kuhn 1939; Romer 1956), Nuthetes destruc-tor (Romer 1956), Poekilopleuron bucklandii (as Mega-losaurus poikilopleuron; Huene 1926a) and Torvosaurustanneri (Paul 1988a). Laelaps trihedrodon Cope, 1877has been previously recombined as Dryptosaurus trihe-drodon (Cope 1878), Creosaurus trigonodon (Osborn1931) and Megalosaurus trihedrodon (Nopcsa 1901). Itis based on a partial dentary (AMNH, lost specimen) andreferred teeth (AMNH 5780) from the Morrison Formation(Kimmeridgian–Tithonian) of Canon City, Colorado thatcan be referred to Allosaurus fragilis (Chure 2001b; Holtzet al. 2004).
Several valid or potentially valid forms are based onspecimens that have at times been assigned to Mega-losaurus but not as distinct species: Becklespinax altispinax(as M. dunkeri), Eustreptospondylus oxoniensis and Strep-tospondylus altdorfensis (both as M. cuvieri). In addi-tion, Dakosaurus maximus Quenstedt, 1858 is a validmetriorhynchoid taxon originally based on a dentary frag-ment from the Kimmeridgian–Tithonian of Schnaitheim,Germany (Megalosaurus sp. in Quenstedt, 1843).
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Other species were erected without a description, diag-nosis or indication of type material and therefore repre-sent nomina nuda (Article 12; ICZN): M. cachuensis fromthe Middle Jurassic Dapuka Group of Xinjiang UygurZizhiqu, China (Zhao in Weishampel et al. 2004b), possiblya misspelling of M. dapukaensis Zhao, 1986 (also a nomennudum; Olshevsky 1991); and Megalosaurus tibetensisZhao, 1986. The remaining species, mostly fragmentaryand non-diagnostic, are assigned as follows and listedalphabetically by species.
Zanclodon cambrensis Newton, 1899. This species isbased 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 toMegalosaurus existed. Waldman (1974) and Molnar (1990)considered that the holotype bore detailed resemblance to‘Megalosaurus’ (now Duriavenator) hesperis and M. buck-landii; Galton (1998, 2005) transferred the taxon to Mega-losaurus cambrensis based on Waldman’s (1974) observa-tions. However, NHMUK R2912 is substantially differentfrom Megalosaurus, indeed from tetanurans generally, andis an indeterminate theropod outside Averostra or a morebasal predatory archosaur (Rauhut & Hungerbuhler 2000;Naish & Martill 2007; Benson 2010b).
Megalosaurus chubutensis del Corro, 1974. M.chubutensis is based on a single lateral tooth (MACN18.189) from the upper member of the Cenomanian(Cerro) Castillo Formation north of Cerro Cretton, Chubut,Argentina (del Corro 1974; Bridge et al. 2000). It ispoorly 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 stronglyrecurved but the posterior is nearly straight. In posteriorview (del Corro 1974, fig. 1) the posterior carina exhibits adistinct apicomedial curvature. The tooth resembles thoseof abelisaurids but is extremely large compared to knownforms.
Megalosaurus cloacinus Quenstedt, 1858. This is amongthe geologically oldest species referred to the genus Mega-losaurus as the holotype and only specimen (SMNS 52457)was found in the Rhat bonebed (Rhaetian–early Hettangian)of Baden-Wurttemburg, Germany. Huene (1932) referredthe holotype dentary of Zanclodon cambrensis to M. cloac-inus (as Gresslyosaurus (?) cloacinus) based on unspeci-fied dental similarity. However, SMNS 52457 is a serrated,recurved tooth of the form typical for theropods. It ismesiodistally slender but does not show any diagnosticfeatures and is therefore Theropoda indet.; additional refer-rals to M. cloacinus cannot be made with confidence.
Megalosaurus dunkeri Dames, 1884. This taxon isfounded on a worn tooth from the ‘lower Wealden’ (EarlyCretaceous) of Nordrhein-Westphalen, Germany (Univer-sity of Marburg N 84; Dames 1884; Huene 1923). Lydekker(1888, pp. 165–168) subsequently referred a large numberof NHMUK specimens from the Lower Cretaceous ofEngland to M. dunkeri although he noted that not all ofthese specimens lacked anterior serrations, a supposedlydiagnostic feature. Lydekker (1890a) later referred ‘mostor all’ of the specimens from Cuckfield, Sussex and the‘Weald Clay’ of the Isle of Wight to Megalosaurus owenifor unspecified reasons. Huene (1923) also considered thatthe holotype vertebrae of Becklespinax altispinax might bereferred to M. dunkeri (q.v.). The holotype of M. dunkeridoes not appear diagnostic beyond the level of Theropodaand referred specimens cannot reliably be assigned to thespecies.
Megalosaurus hungaricus Nopcsa, 1902. Nopcsa (1902,figs 1–6) described and illustrated a single tooth (MAFIOb 3106) from the Late Cretaceous coal-bearing strataof Nagy-Baroth, Romania (then part of the Austrian-Hungarian Empire) as the holotype of M. hungaricus.Nopcsa considered this stratum equivalent to the Maas-trichtian beds at Szentpeterfalva but the proximity of thislocality to the Early Cretaceous Brusturi site suggestscaution in the absence of detailed geological reconnais-sance. 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 byspaces equivalent to the proximodistal width of a singledenticle (Nopcsa 1902, p. 104). The anterior denticles areslightly smaller than the posterior ones, with a denticlesize difference index around 1.25 (Nopcsa 1902), simi-lar to those of some tyrannosauroids and dromaeosaurids(Rauhut & Werner 1995; Sweetman 2004). M. hungaricusteeth are less transversely compressed than those from thesame general age in Europe referred to M. pannoniensis(Nopcsa 1902). Nonetheless, the latter has been consid-ered a senior synonym of M. hungaricus (Lapparent 1947;Lapparent & Zbyszewski 1957). Huene (1926a, b) notedthat ‘M.’ hungaricus could not be assigned to a genus andwe concur that the taxon should be considered Theropodaindet., although it is possible that they represent either tyran-nosauroids or dromaeosaurids.
Megalosaurus inexpectatus del Corro, 1966. This secondspecies of Megalosaurus from the Chubut Group wasbased on five large teeth (holotype, MACN 18.172) asso-ciated with the holotype skeleton of the sauropod Chubuti-saurus insignis (del Corro 1975). They derived from theAlbian–Cenomanian Bayo Overo Member of the CerroBarcino Formation at Paso de Indios, Argentina (del Corro1966). At least two of the teeth (del Corro 1966, pl. 1, figs1, 2) exhibit distinct banding of the enamel as is common
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among tetanurans (Brusatte et al. 2007). The holotypeshows an overall curvature and apical wear reminiscent ofcarcharodontosaurid teeth, to which it should be compared.
Megalosaurus ingens Janensch, 1920. This taxonconsists of a large tooth (MB.R1050; 15 cm long) fromTendaguru Quarry B in the Upper Dinosaur Member ofTanzania (Tithonian; Janensch 1920, 1925). Both anteriorand posterior carinae are coarsely serrated (≤1 per mm)along their entire lengths. The enamel shows transversebands that are particularly pronounced posteriorly as inmany tetanurans (Brusatte et al. 2007). Numerous otherteeth (25 MB specimens; NHMUK R5556, R5557, R6758),some from other levels of the Tendaguru Beds, have beenreferred to this taxon based primarily on the coarseness ofthe serrations and their large overall size (Janensch 1925).We do not consider these characteristics sufficient to definea taxon or refer additional specimens, and refer M. ingensto Tetanurae indet.
Megalosaurus insignis Eudes-Deslongchamps &Lennier in Lennier, 1870. The holotype tooth of M.insignis was found in the lower Kimmeridgian depositsof Cap de la Heve, France (Valenciennes 1863; Lennier1870; Huene 1932). It was described as large (80 mmlong), laterally compressed, recurved, and serrated alongthe posterior carina but was missing its apex (Lennier1870, 1887, pl. 1, figs 1–3). It is indeterminate belowthe level of Theropoda. Additional specimens include apedal phalanx and ungual from the same area (Lennier1887; 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 ofPortugal were later assigned to M. insignis (Lapparent& Zbyszewski 1957); although some elements preserveinteresting morphological details, placement within thistaxon is not justified. The Portuguese collection includesan articulated series of five anterior caudal vertebraefrom Praia da Areia Branca that are associated with twoadditional articulated caudals. The transverse processesare upturned and extend far laterally whereas the centralack neural arch laminae or centrum pleurocoelous fossaeand bear tall, vertical neural spines. They may belong to ateleosaurian crocodilian (Chabli 1986).
Megalosaurus lonzeensis Dollo, 1883. This species wasfounded on a partial manual ungual from the Santonian‘glauconite argileux’ of Lonzee, Namur, Belgium (Dollo1883). The small size, mediolaterally narrow dimensionsand details of the vascular traces more closely resemblethe manual unguals of coelurosaurs. Huene (1926a, 1932)placed it among ornithomimids but it is best referred to asCoelurosauria indet.
Megalosaurus lydekkeri Huene, 1926a. A tooth from theLower Lias Group of Lyme Regis (NHMUK 41352) maycorrespond to reports of Megalosaurus (Dawkins in Huxley1870) or M. bucklandii (Phillips 1871) from the Lias ofDorset. It was first formally recorded as Zanclodon sp. b(Lydekker 1888) and later questionably assigned to Mega-losaurus sp. (Huene 1908), ‘Zanclodon’(?) sp. (Huene1926a, p. 36) and ‘Megalosaurus’ (gen.?) lydekkeri n. sp.(Huene 1926a, table 1). The crown is laterally compressedand smoothly curved throughout its length, bearing nearlyfully serrated anterior and posterior carina and faint longi-tudinal striations. The latter may suggest non-tetanuranaffinities, because the tooth lacks the specialized featuresof tetanurans in which such striations also occur (e.g.spinosaurids). Otherwise the specimen is indeterminatewithin Theropoda.
Megalosaurus meriani Greppin, 1870. This fragmen-tary form derives from the ‘Virgulian’ Basse Montagnequarry near Moutier, Kanton Bern, Switzerland, now knownto belong to the lower Reuchenette Formation (earlyKimmeridgian). It was originally described as ‘a completeskeleton’, ‘a large part’ of which was in the Musee de Bale(now the Naturhistorisches Museum Basel). However, theaccompanying description and plate (Greppin 1870, pl. 1,figs 2–5) demonstrate that the majority of the specimenspertain to a sauropod; these were later removed from M.meriani and now constitute the holotype of Cetiosauriscusgreppini (Huene 1927; Meyer & Thuring 2003). The holo-type of M. meriani is therefore restricted to the only demon-strably theropod element, a single tooth (MH 350; Greppin1870, pl. 1, fig. 1). It is mediolaterally thick, bears longi-tudinal striations on the internal face and fine serrationson both carinae. Overall it resembles the anterior teeth ofCeratosaurus (Madsen & Welles 2000), although we assignit to Ceratosauria indet. rather than this genus.
Megalosaurus mersensis Lapparent, 1955. M. mersen-sis was based on a partially articulated series of 23 verte-brae from the Bathonian El Mers Formation north of Tizin’Juillerh, Morocco (Lapparent 1955). These are not wellpreserved but represent much of the neck and back as wellas parts of the sacrum and tail. The axis lacks pneumaticforamina and bears only a small diapophysis. It is diffi-cult to determine from the illustrated vertebrae (Lapparent1955, pl. 3) whether any of the subsequent vertebral centraare pneumatized. The cervical centra appear to have onlyweakly offset faces and are amphicoelous (Chabli 1986).The anterior dorsal (Lapparent 1955, pl. 3, fig. 6) has ahorizontally oriented transverse process and a neural spineat least as tall as the height of the centrum. The centra of twosucceeding dorsals (Lapparent 1955, pl. 3, fig. 8) are slightlylonger than tall and have a shallow ventral arch. The verte-brae are quite different from those of most megalosauroids(e.g. Baryonyx, Eustreptospondylus, Piatnitzkysaurus) in
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one or more of these features. Without additional study ofthe type specimen we cannot determine the validity of thespecies ‘M.’ mersensis, although we agree that it does notbelong within the genus Megalosaurus and instead seemslikely to represent a teleosaurid mesosuchian (Chabli 1986).
Saurocephalus monasterii Munster, 1836. Munster(1836, pl. 3, fig. 15a–d) erected a new species of the fishgenus Saurocephalus, S. monasterii based on a recurved,serrated tooth from the Oxfordian Korallenkalk of the Lind-ner Berge, Hanover, Germany. Windolf (1997) transferredthe taxon to Megalosaurus but Munster’s tooth cannot beidentified past the level of Theropoda indet.
Aggiosaurus nicaeensis Ambayrac, 1913b. A partialmandible from the upper Oxfordian in the region of Capd’Aggio-La Turbie, Monaco was reported as a plesiosaur orteleosaurid crocodile by Deperet (in Ambayrac 1913a) andmade the holotype of the crocodilian taxon Aggiosaurusnicaeensis (Ambayrac 1913b). Subsequently the materialwas considered to represent a carnosaurian (Huene 1932)and transferred to Megalosaurus nicaeensis (Romer 1956;Steel 1970). The teeth of A. nicaeensis are conical andonly slightly recurved, with fine enamel ridges; replace-ment teeth are present inside the hollow roots of functionalteeth. Therefore, A. nicaeensis can be placed within Metri-orhynchidae, although as a nomen dubium (Buffetaut 1982).
Megalosaurus obtusus Henry, 1876. The holotype spec-imens of this form were recovered from Rhaetian marineoutcrops near Moissey, Franche-Comte, France. The fourreferred teeth are fragmented and poorly preserved but showdistinct carinae and serrations (Henry 1876, pl. 1, figs 1–4).Thus they should not be referred to Plateosaurus (Huene1908) but as indeterminate specimens within Theropoda oranother predatory archosaur clade.
Megalosaurus pannoniensis Seeley, 1881. The Maas-trichtian Gosau Formation near Weiner Neustadt, Austriaproduced two teeth (GMUV uncatalogued) that Seeley(1881) referred to M. pannoniensis (for which M. pannon-icus is a lapsus calami; Huene 1926a). Huene (1926a)considered them unlikely to pertain to Megalosaurus dueto their young geological age and noted that the illustratedspecimen “does not show anything concerning the natureof the genus” (Huene 1926a, p. 81). The serrations are fine,∼ 3.5 per mm on the anterior carina and ∼ 2 per mm on theposterior carina. The anterior serrations terminate beforethe base of the tooth and Seeley (1881, p. 670) noted thepresence of transverse “lines of growth” across the enamelthat are now regarded as enamel ‘wrinkles.’ Together thesefeatures allow assignment to Tetanurae indet., and the rela-tively small anterior denticles suggest possible assignment
to either Dromaesauridae or Tyrannosauroidea (Rauhut &Werner 1995; Sweetman 2004).
Megalosaurus pombali Lapparent & Zbyszewski, 1957.This species was named for three teeth from unnamedCallovian–Oxfordian beds near Pombal, Portugal. Lappar-ent & Zbyszewski (1957) distinguished it from M. insignisby 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 shouldbe relegated to Theropoda indet. However, a set of eightvertebrae from Porto das Barcas was also referred to M.pombali. Although poorly described and figured and notreferable to M. pombali, they warrant further study to deter-mine whether they represent a diagnosable taxon.
Massospondylus rawesi Lydekker, 1890b. The holotypetooth, from the Takli Formation (late Maastrichtian; Sahniet al. 1984) of Maharashtra, India, is recurved, serratedand laterally compressed (NHMUK R4190). Although thespecies has been referred to Megalosaurus (Vianey Liaudet al. 1987) and Orthogoniosaurus (Olshevsky 1991), it isnot diagnostic past the level of Theropoda indet. The fineserrations and stout proportions resemble the condition inabelisaurids (Carrano et al. 2010).
Megalosaurus schnaitheimii Bunzel, 1871. This taxonis based on teeth originally described and illustrated asMegalosaurus sp. by Quenstedt (1852). They derive fromLate Jurassic marine deposits (possibly the Korallenkalk)of Schnaitheim, near Brenz, Germany, and may pertain toDakosaurus maximus.
Zanclodon silesiacus Jaekel, 1910. A tooth from theMuschelkalk of Gogolin, Oberschlesien, in the collectionof the University of Griefswalden/Gottinger is the holotypeof Zanclodon silesiacus Jaekel, 1910. This specimen wasreferred to ‘Megalosaurus’ silesiacus by Kuhn (1965). Ithas the recurved, serrated form typical of theropods andcould be considered as Theropoda indet., but we cannotrule out the possibility that it represents a ‘rauisuchian’archosaur.
Megalosaurus terquemi Gervais, 1859. This species isbased on three isolated teeth from the Hettangian ‘Angula-tus Beds’ of Hettingen, near Moselle, Germany (Gervais1853; Terquem 1855; Huene 1926a). Huene consideredthem sufficiently distinct to refer to them as ‘Mega-losaurus’ (gen. 2) terquemi (Huene 1926a, p. 80) orMegalosauridorum gen. indet. terquemi (Huene 1932 p.219), noting that the teeth were longer, sharper andmore lingually compressed than those of Plateosaurus(Huene 1932). Buffetaut et al. (1991b) considered thatthe teeth belonged to phytosaurs. As they were housed
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at the Museum du Caen, we presume they were lostwhen that institution was destroyed by incendiary bombsduring the Second World War, although they were notspecifically listed as such (Bigot 1945). We refer them toArchosauria indet.
Finally, the tendency to assign tracks to Megalosauridaeor Megalosaurus is largely without a firm anatomical basis,and all such occurrences should be considered Theropodaindet. This is particularly true of large theropod trackwaysfrom the Early Cretaceous of England (Delair 1982) andSpain (Sanz et al. 1990), which are far removed temporallyfrom any European occurrences of this family.
It is beyond the scope of this paper to review all theindividual specimens assigned to the genus Megalosaurus,but we are doubtful that any (aside from those specificallyreviewed above and in Benson 2010a) have been correctlyattributed.
Fragmentary occurrencesAfrica. The Late Jurassic–?earliest Cretaceous TendaguruFormation includes a small number of theropod specimenscompared to the penecontemporaneous Morrison depositsof North America, but probably includes 4–5 taxa nonethe-less. Among them are a slender ceratosaur, Elaphrosaurus;a larger, more robust relative that includes some of the mate-rial referred to Ceratosaurus? roechlingi and several hindlimb bones (Janensch 1925; Carrano et al. 2002; Carrano &Sampson 2008); and at least one small abelisauroid (Rauhut2005b). 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 rightilium (MB.R 3628 = St 233) also derives from a tetanu-ran (Fig. 11) but is unusual in exhibiting a very widelyopen preacetabular notch, a strongly but broadly down-curved dorsal margin of the preacetabulum, and a markedlytapering 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 pedun-cle with length:width proportions similar to allosaurians(∼ 2.0) but shorter than coelurosaurs (∼ 3.0) and longerthan all non-avetheropods and metriacanthosaurids (≤1.7),and a ventrally rather than anteroventrally oriented pubicpeduncle as in carcharodontosaurians. Taken together, thesefeatures suggest that the ilium may pertain to a carchar-odontosaurian. The many isolated Tendaguru teeth arelargely Theropoda indet., although some may pertain toceratosaurians as they possess longitudinal striations on thelingual surface similar to those on the premaxillary teeth ofCeratosaurus (Madsen & Welles 2000).
Approximately contemporaneous with the Tendagurumaterials, theropod teeth from the Late Jurassic ofEthiopia have been described as cf. Acrocanthosaurus sp.(Goodwin et al. 1999). However, this identification wasbased solely on the presence of ‘chisel-shaped’ denticles, afeature present in, but not unique to, Acrocanthosaurus.One specimen displays surface crenulations (‘wrinkledenamel’), but these are widely distributed among tetanurans(Brusatte et al. 2007) and we assign these teeth to Tetanuraeindet.
The more southerly exposures of the ?Late JurassicKadzi Formation of Zimbabwe have also produced thero-pod remains, including femora referred to Allosauridae?indet. (Raath & McIntosh 1987). These femora are badlydamaged but exhibit several features that suggest affinitieswith Ceratosauria, rather than Tetanurae. They are currentlyunder study (E. Roberts, P. O’Connor & M. Carrano inprep.).
A vertebral centrum from the Hauterivian–BarremianCabao Formation of Libya was referred to ?Spinosaurussp. (El-Zouki 1980; Tawadros 2001). The proportionsmatch well with the posterior dorsals of Baryonyx andSuchomimus but are relatively shorter than those of Mega-losaurus and carcharodontosaurids. The (presumed) ventralsurface is only slightly narrower than the centrum, simi-lar to the condition in Megalosaurus and Suchomimus(El-Zouki 1980, pl. 2). We consider it likely thatthis specimen belongs to the clade Megalosauria. Thetooth listed as cf. Baryonyx sp. from the same forma-tion (Le Loeuff et al. 2010) is assigned here toBaryonychinae indet.
A proximal femur (AM 6041) from theBerriasian–Valanginian Kirkwood Formation (Forster etal. 2009) of South Africa indicates the presence of a smalltetanuran in these deposits. Forster et al. (2009) proposedthat the inferred presence of a weakly anteromedially(rather than strictly medially) oriented femoral headsuggested non-avetheropodan affinities. However, thefemoral head of some allosauroids is oriented slightly(Allosaurus, Sinraptor; Benson 2009b, 2010a) or strongly(Neovenator; Brusatte et al. 2008) anteromedially andboth the absence of a proximal articular groove andthe presence of a well-developed accessory trochanter
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indicate avetheropodan affinities. We regard this specimenas Allosauroidea indet. but note that the head lacks theproximal inclination of most carcharodontosaurians.
Spinosaurid and carcharodontosaurid remains, mostlyteeth but also vertebrae (Rauhut 1999), have been foundacross North Africa in deposits ranging in age from Albianthrough Cenomanian (e.g. Bouaziz et al. 1988; Russell1996; Rauhut 1999; Sereno et al. 2004). It is not clearwhether all these specimens can be accommodated bycurrently named taxa, but this issue is unlikely to be decidedwithout more complete materials.
Sereno & Brusatte (2008) referred postcranial spec-imens including a pelvis and sacrum (MNN GAD1-2)from the Elrhaz Formation (Aptian–Albian) of Niger to theabelisaurid Kryptops palaios. This material was collected insitu but the holotypic maxilla (MNN GAD1-1) was founderoded free of the rock some 15 metres distant. Severalfeatures of the pelvis and sacrum were described as ‘strik-ingly primitive’ for an abelisaurid, such as the presence offive sacral vertebrae, a reduced supraacetabular shelf thatis not continuous with the lateral wall of the brevis fossa,an iliac pubic peduncle that is substantially larger than theischial peduncle, and the relatively high and dorsoventrallyshort proportions of the ilium (Sereno & Brusatte 2008,p. 23). However, these features, which are present gener-ally in tetanurans, are not present in even the most basalceratosaurs and would therefore represent homoplasies inan abelisaurid. Both Ceratosaurus and Elaphrosaurus havesix sacral vertebrae, a hypertrophied supraacetabular shelfthat is continuous with the lateral wall of the brevis fossa,an iliac pubic peduncle that is only slightly larger than theischial peduncle, and relatively long, low proportions ofthe iliac blade. Furthermore, the ischia of MNN GAD1-2contact via a medial ‘apron’ as in many tetanurans butunlike the flat medial contact of more basal theropods,whereas the presence of a ventral fenestra between the pubicdistal expansions similar to that of Ceratosaurus (Bensonet al. 2009), as figured by Sereno & Brusatte (2008, fig.7B), cannot be determined due to damage. Instead, a prox-imodistally elongate fenestra is present between the pubicshafts proximal to the distal expansion, as seen in tetanu-rans (Benson et al. 2009). The presence of a subrectan-gular fenestra between the fourth and fifth sacral neuralspines (Sereno & Brusatte, 2008, fig. 7) is identical to thatof Giganotosaurus (MUCPv-Ch 1), as is the presence ofa peg-and-socket iliac-ischial articulation (a carcharodon-tosaurian synapomorphy). Taken together, we consider itunlikely that MNN GAD1-2 pertains to an abelisaurid oreven a primitive ceratosaur, and suggest that is representsa carcharodontosaurid, most likely the sympatric Eocar-charia (currently only known from skull bones).
Asia. Nessov (1995) described ‘coelurid’ and ‘mega-losaurid’ teeth from the Callovian Balabansai Svita atSarykamyshai in the Fergana Valley, Kyrgyzstan. These
were recently redescribed (Averianov et al. 2005) as belong-ing to tetanurans with possible affinities to Dromaeosauri-dae. The presence of a short anterior carina was noted asa particular similarity with dromaeosaurids, but in fact thisfeature is quite common among basal tetanurans and isrecovered here as the primitive condition for the clade.Other aspects of the size and shape of these teeth are alsoconsistent with identification as a non-coelurosaurian teta-nuran.
Six teeth from the Callovian–Oxfordian DjaskoianFormation of Teete Creek, Yakutia, Russia were identi-fied as Allosaurus sp. (PIN 4874/2; Kurzanov et al. 2003).They are large, recurved and serrated along both carinabut cannot be identified more specifically than Theropoda.Likewise, Allosaurus sp. has been reported from unspec-ified Late Jurassic strata near Datong, in the XinrongDistrict of Shaanxi Province, China (Lu & Hu 1998). Theyinclude anteroposteriorly short dorsal vertebrae that appearto bear deep lateral fossae, ‘honeycomb’ internal structuredescribed by Lu & Hu (1998, pl. 1) seems to be a marrowcavity rather than camellate pneumatic architecture. Theoverall morphology and cavernous lateral depressions areloosely comparable to Torvosaurus posterior dorsal verte-brae. However, the published images are difficult to inter-pret, so we consider them Tetanurae indet cf. Torvosaurus.
Buffetaut & Suteethorn (2007) reported a metriacan-thosaurid (‘sinraptorid’) tibia from the Late Jurassic–EarlyCretaceous Phu Kradung Formation of Thailand. Asreported by these authors, this specimen (SM 10) showsvarious detailed similarities with Chinese metriacan-thosaurid taxa that support this identification.
Early Cretaceous deposits at Tsanmakou in Shaanxi,China have yielded a fragmentary theropod skeleton (IVPPV969) that was assigned to Allosauridae (Young 1958).The material was only briefly described and illustratedbut was compared favourably to Antrodemus valens (=Allosaurus fragilis); Young (1958, p. 235) suggested itmight represent a new taxon. The scapula was describedas being robust and having a somewhat expanded distalend. The fibula differs from those of spinosaurids and othermegalosaurians in possessing a prominent, centrally placedmedial fossa, and from those of allosauroids in exhibiting anobliquely oriented, rather than horizontal, proximal marginof this fossa. Unlike tetanurans generally, the iliofibularistubercle is quite prominent. This material may represent aceratosaurian but further study is required. We refer it toAverostra 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 possi-ble carcharodontosaurid, although they noted several differ-ences with the teeth of currently known forms. Both thearcuate wrinkles and overall proportions are reminiscent ofcarcharodontosaurids but are now known to occur widelywithin Tetanurae (Brusatte et al. 2007).
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A middle caudal vertebra, fragmentary tibia and the distalend of a phalanx (IVPP V2756) from the Early Creta-ceous Jehol Group of Shaihaitzu, Liaoning were assignedto Megalosauridae (Hu 1963). However, the tibia (theonly illustrated element) bears a small, discretely roundedlateral condyle at the proximal end that strongly recalls themorphology of more derived theropods. It may pertain to acoelurosaurian.
A fragmentary tibia from the Upper Cretaceous of Syria(Hooijer 1968) probably represents a tetanuran based onits described similarity to the same bones in Carcharodon-tosaurus and Erectopus. In particular, the proportions ofthe distal end are transversely expanded, comparable to thecondition in tetanurans generally.
Other theropod teeth from Jurassic and Cretaceousdeposits of eastern Asia, often ascribed to megalosaurids(YNUGI 10003; KS 7001; Hasegawa et al. 1992; Nessov1995; Lim et al. 2002) or allosaurids (KPE 8004, 8005; Parket al. 2000) cannot be diagnosed beyond Theropoda (e.g.Averianov et al. 2005). Those from South Asia (Ghevariya1988; Sahni & Bajpai 1988) are similarly indeterminate(e.g. Bajpai & Prasad 2000), although many may ultimatelypertain to abelisaurids.
Australia and Antarctica. Aside from Cryolophosaurusand Australovenator, most theropod remains from thesecontinents are quite fragmentary. The purported tetanu-ran caudal vertebra (WAM 96.5.1; Long & Cruickshank1996) from the Birdrong Sandstone north of Kalbarri,Western Australia is too fragmentary to identify witheven this level of specificity, and we consider it to beTheropoda indet. We likewise assign the ‘carnosaur’ pedalphalanx from the Cenomanian–Turonian Molecap Green-sand (WAM 92.7.1; Long 1995) and the ungual from theValanginian–Albian Wonthaggi Formation once referredto Megalosaurus (NMV P.10058; Woodward 1906; Huene1926a; Rich & Rich 1989) to Theropoda indet.
A theropod ulna (NMV P186076; Rich & Vickers-Rich2003) from the Eumeralla Formation at Dinosaur Cove,Victoria (Aptian–Albian) was referred to cf. Megaraptor bySmith 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 theConiacian–Santonian Hidden Lake Formation of JamesRoss Island was identified as Tetanurae indet. (Molnaret al. 1996). These authors noted similarities with basaltetanurans, particularly Piatnitzkysaurus, in the shapes ofthe medial malleolus, medial buttress and fibular flange.The distal end has a ratio of transverse width to anteropos-terior 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 hasbeen reduced by abrasion in MLP 89-XII-1-1. The boneis 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, sowe suggest that this bone pertains to a megalosauroid, thefirst identified from either continent, and one of the latestknown.
Perhaps the best known tetanuran specimen fromAustralia is an astragalus recovered from the WonthaggiFormation at Eagle’s Nest, Victoria (NMV P150070)that was originally referred to Allosaurus sp. (Molnaret al. 1981). Since its description, its taxonomic iden-tity has been much debated (Welles 1983; Molnar et al.1985; Rich 1996; Chure 1998) and it was most recentlyassigned to Australovenator sp. (Hocknull et al. 2009) andAbelisauroidea (Agnolin et al. 2010). Despite the argu-ments of Agnolin. et al. (2010), we agree that this speci-men pertains to a megaraptoran allosauroid, but given itstemporal and geographical distance from the type materialof Australovenator, and its resemblance to the astragalus ofFukuiraptor, we cannot exclude that it belongs to a distinctmember of this clade (Benson et al. 2010).
Europe. The theropod knee joint once considered partof the lectotype of Scelidosaurus harrisonii (NHMUK39496; Newman 1968) shows several tetanuran character-istics (Carrano & Sampson 2004). The proximal morphol-ogy of the tibia resembles the condition in megalosauroidsmore so than ceratosaurs or coelophysoids. The fibula hasa large but shallow, posteriorly open fossa on the medialsurface as in some basal tetanurans and non-megalosaurianmegalosauroids, but unlike the deep, centrally placed fossaof ceratosaurs or the narrow posterior incisure of coelo-physoids. Finally, the distal femur has a vertically orientedtibiofibularis crest as in most tetanurans. This Liassic spec-imen may therefore be one of the earliest known tetanu-rans, but until more complete remains are found it is uncer-tain whether these features are synapomorphies permittinginclusion within Tetanurae, or symplesiomorphies that aretransformed in Ceratosauria (Benson 2010b).
The informal name ‘Saltriosaurus’ has been used torefer to a large theropod discovered in the SinemurianSaltrio Formation, near Saltrio, Varese, Italy, in 1996(Dal Sasso & Brillante 2001; Dal Sasso 2003). Thespecimen (MSNM V3664) is incomplete but includes afurcula, scapula, humerus, manual elements and parts ofthe hind limb. Enough is preserved to support the theropodaffinities of ‘Saltriosaurus,’ but more specific assignmentmust await further preparation and description of thematerial. The widespread presence of a furcula amongsttheropods, including coelophysoids (Tykoski et al. 2002;Rauhut 2003; Rinehart et al. 2007), demonstrates that thisfeature alone is insufficient to identify ‘Saltriosaurus’ as atetanuran (Benson 2010b).
Little can be said of the theropod vertebra describedas Megalosaurus by Huene (1966) from a glacial erratic
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boulder in Ahrensburg, Schleswig-Holstein, Germany,which he considered to have been derived from the ‘Liasepsilon’ (Toarcian). Interestingly, the fragmentary speci-men exhibits a convex anterior centrum face and a substan-tial lateral pleurocoelous fossa. It may therefore representa 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 theropodbones from the early Bathonian of New Park and OakhamQuarries to Megalosaurus. Benson (2009b) reviewed theNew Park Quarry material and established that all boneswere either non-diagnostic or referable to Megalosaurus,with no evidence for a second theropod taxon. The anatomyof these bones cannot be distinguished from M. bucklandiiat present. The material from Oakham Quarry is currentlyunder review (Benson 2009a), but some of the bonesshow autapomorphies of Megalosaurus whereas others (e.g.SDM 44.19, a partial ilium) represent a basal tetanuran ofuncertain affinities that is distinct from either Megalosaurusor Cruxicheiros. At least 39 other localities in the BritishBajocian–Bathonian have yielded isolated theropod bonesand teeth, most of which are indeterminate (Benson 2009a).In summary, three large bodied basal tetanurans, Mega-losaurus, Cruxicheiros, and an additional unnamed taxonare known from the early–middle Bathonian of England(Fig. 12).
The Callovian Vaches Noires cliffs of northern Francehave produced several fragmentary theropod specimens inaddition to the holotype of Piveteausaurus. Among themis a partial braincase from Auberville (Bulow collection25192; Knoll et al. 1999) that probably pertains to amegalosauroid, although referral to Allosauroidea cannotbe excluded. A pair of articulated frontals from Houl-gate (Enos collection; Buffetaut & Enos 1992) can bereferred to Tetanurae based on the presence of a distinctsocket for articulation with the prefrontal. The anteroposte-riorly long dimensions of these frontals suggest placementamong megalosauroids or other basal tetanurans rather thanavetheropods. Knoll et al. (1999, p. 106) suggested that thespecimen might pertain to the same taxon as the brain-case, which would further imply that the latter is a mega-losauroid. Both elements are very similar to the holotypeof Piveteausaurus, especially given our reinterpretation ofthe sutural contacts in the latter, and we suggest that theymay be referable to this form. Finally, a maxilla fragmentfrom Villers-sur-Mer was referred to Megalosaurus sp.(Pennetier collection 380; Buffetaut et al. 1991a). It bearslarge foramina on the lateral surface and gaps between theroughly surfaced paradental plates that are more consistentwith 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 mega-losauroids among contemporaneous French and Englishspecimens, a long snouted form (Eustreptospondylus) and
a shorter snouted form. The second taxon is represented byat least the maxilla, which has a short anterior process, butmight also include the holotype of Piveteausaurus (as wellas the isolated frontal and braincase).
‘Megalosaurid’ specimens from the Late OxfordianSables de Glos of Lisieux, Basse-Normandie, France(Buffetaut et al. 1985) comprise only serrated, laterallycompressed teeth that should be regarded as Theropodaindet.
Theropod remains are rare in theKimmeridgian–Tithonian deposits of Europe, knownprimarily from Portugal (Allosaurus, Aviatyrannis,Ceratosaurus, Torvosaurus) and England (Stokesosaurusand various ‘megalosaur’ remains). Powell (1987, p. 108)referred a worn maxilla fragment from a large theropod thathad been dredged from the seabed west of Portland, Dorset(DORCM G10603; Aulacostephanus autissiodorensisAmmonite Zone; lower Kimmeridgian) to Megalosauridae.The presence of an anteroventrally inclined anterior sectionof dorsal boundary of the paradental plates supportsthis assignment, and the dorsoventral striations on theparadental plates may allow identification as Megalosaurus(Benson & Barrett 2009). We consider the specimen asMegalosauridae indet., cf. Megalosaurus, and note that italso shares very tall paradental plates with Megalosaurusand Torvosaurus. Fragmentary large theropod remainscomprising a right tibia (OUMNH J.29886; Huene,1926a) and left metatarsus (OUMNH J.13586a–c; Phillips1871) were also recovered from unknown horizons ofthe Kimmeridge Clay Formation (Kimmeridgian–earlyTithonian) at Swindon, Wiltshire. The half-century intervalbetween these reports suggests that they were not found inassociation. All were referred to Megalosaurus bucklandii(Phillips 1871; Huene 1926a; Delair 1973), althoughHuene (1932) later revised this to Megalosaurus (?)sp. The proportions of the tibia are similar to those ofTorvosaurus, the most robust basal tetanuran, but there islittle else to allow firm identification and they are regardedhere as Tetanurae indet. These specimens are consistentwith the presence of a robust megalosaurid in the UK,contemporaneous (and possibly conspecific) with thePortuguese Torvosaurus.
Many other theropod elements from the KimmeridgeClay Formation should be regarded as Theropoda indet.,including phalanges from the lower Kimmeridgian Rase-nia cymodoce Zone of Wyke Regis, Dorset (Broken-shire & Clarke 1993; Benson & Barrett 2009), a toothfrom Foxhangers, Wiltshire referred to Megalosaurusinsignis (NHMUK 46388; Lydekker 1888), ‘megalosaur’phalanges from an unspecified locality in Wiltshire(DZSWS 3009; Delair 1973), and a proximal caudal verte-bra from Shotover, Oxfordshire (OUMNH J.47134; Powell1987).
The fragmentary pectoral remains of a large theropodfrom the early Valanginian of Montmirat, France were
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Figure 12. Stratigraphical relationships of Jurassic tetanurans in England and France. The central columns show the approximatestratigraphic 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-coelurosaurianavetheropods; stars, coelurosaurs.
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described by Perez-Moreno et al. (1994) and interpretedas a close relative of Allosaurus. The specimen, whichincluded a partial scapula, humerus and manus, displaysproportions that are similar to those of Allosaurus but alsoother basal tetanurans (e.g. Afrovenator). Three charac-ters were used to link it with Allosaurus (humerus/scapulalength ratio <65%; metacarpal I strongly curved medially;metatarsal III/tibia length ratio <54%) but do not supportsuch a grouping here. We capture the first two morpholo-gies with different character descriptions and codings; thethird is heavily influenced by scaling and locomotor habitand shows no clear phylogenetic signal. We suggest thatthis form be referred to as Tetanurae indet.
Numerous spinosaurid teeth as well as occasionalskeletal elements from the Early Cretaceous of Spain,Portugal and England have been assigned to Spinosauri-dae, Baryonychinae or Baryonyx (e.g. Viera & Torres1995; Martill & Hutt 1996; Torcida et al. 1997; Ruiz-Omenaca & Canudo 2001; Canudo & Ruiz-Omenaca2003; Pereda Suberbiola et al. 2003; Canudo et al.2004a, 2008; Ruiz-Omenaca et al. 2004; Infante et al.2005; Torcida Fernandez-Baldor et al. 2006; Gomez-Fernandez et al. 2007a, b), although many have yet to bedescribed or illustrated. Certainly some of these pertain tobaryonychines, particularly a partial spinosaurid maxilla(GA-2065) from the Enciso Group of Igea (Viera &Torres 1995) that lacks the paired alveoli characteristicof spinosaurines. Currently, only a partial skeleton (ML1190) from the Barremian of Portugal is complete enoughto warrant assignment to Baryonyx sp. (Mateus et al.2011).
The presence of a carcharodontosaurid in the MorellaFormation (early Aptian) has been suggested based on asingle distinctive tooth (Canudo & Ruiz-Omenaca 2003).Benson et al. (2009) recently reported a partial pubis andfemur (MIWG 6350) from the Wessex Formation (Berri-asian) of the Isle of Wight that indicated a basal tetanurandistinct from the sympatric Baryonyx and Neovenator. Thisspecimen lacks autapomorphies but probably represents anallosauroid rather than a more basal tetanuran based on theenlarged (but abraded) distal pubic expansion. Naish (2003)also reported the proximal end of an allosauroid tibia fromthe stratigraphically higher Hastings Beds Group (upperBerriasian–Valanginian). It cannot be established whetherthis represents a distinct taxon from the incompletely knowncoeval forms Becklespinax and Valdoraptor.
Both megalosauroids and carcharodontosaurians areunknown in any Late Cretaceous deposits from Europe,which have yielded only a few fragmentary remainsof large-bodied theropods. Supposed megalosaurid teethfrom the Maastrichtian of Aveiro and Taveiro, Portu-gal (CEPUNL AV1–2, TV 36, 41, 42, 49; Antunes &Sigogneau-Russell 1991, 1992) can only be assigned toTetanurae (although they are more slender and finelyserrated than tyrannosaurid teeth).
Casanovas-Cladellas et al. (1988) described an isolatedradius (IPSN-18) from Els Nerets in the Late Cretaceous(Maastrichtian) Tremp Group of Spain as belonging toMegalosauridae. However, this specimen is not diagnosticto the family level and should be considered an indetermi-nate large theropod (Canudo & Ruiz-Omenaca 2003).
North America. Aside from Acrocanthosaurus, putativecarcharodontosaurid teeth are known from the Early Creta-ceous Patuxent Formation (Arundel Clay facies, Aptian)of Maryland. These include several specimens assignedto Acrocanthosaurus (Lipka 1998) but which, aside frombeing penecontemporaneous with this taxon, show nodiagnostic features below the family level. A large toothcollected from Coffin’s Old Engine Bank, near Muirkirk,was originally housed at Goucher College (#3121) andsupposedly donated to the USNM, but no record of itsarrival exists; a published photograph is the only evidenceof this unusually fine Arundel theropod specimen (asAllosaurus; Bibbins 1895). In all cases, the teeth resemblethose of carcharodontosaurians but do not possess unequiv-ocal synapomorphies and can only be assigned to Tetanuraeindet.
Several ‘?megalosaurid’ specimens were described fromthe Aptian–Albian Cloverly Formation of Wyoming andMontana (Ostrom 1970). Many were teeth (YPM 5369,5377–5379) that can be considered Theropoda indet. Ametatarsal III (YPM 4885) bears features common to manytheropods, although it does seem to differ from mostcoelurosaurs. A posterior dorsal vertebra (YPM 5285) isanteroposteriorly short but differs from those of Acrocan-thosaurus in the relatively short neural spine and lackof centrum pneumaticity (Ostrom 1970; Harris 1998b).Like Becklespinax, this material can be excluded fromSpinosauridae and Carcharodontosauria but not placedmore specifically within Tetanurae.
Allosauroids and megalosauroids are currently unknownfrom the early Late Cretaceous of North America, althoughit is poorly sampled prior to the Campanian. The very well-sampled latest Cretaceous has also yielded no evidence ofeither clade.
South America. In addition to the named materials fromthe Canadon Asfalto Formation, Rauhut (2007) described apartial theropod skull (MPEF-PV 1717) that differed fromthe contemporaneous Piatnitzkysaurus. Rauhut (2007)surmised that it might belong to either Condorraptor (forwhich no skull materials are known) or a third, unknowntaxon. Regardless, the specimen clearly represents at leasta ‘stem’-tetanuran, as shown by the presence of a distinctmaxillary fenestra (Rauhut 2007).
Additional tetanuran remains from South Americacomprise a series of dorsal vertebrae associated with apartial manus (SNGM-1887, 1894, 1898, 1900, 1903) fromthe Tithonian Toqui Formation of southern Chile (Salgado
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et al. 2008). The material shows tetanuran features (fuseddistal carpals 1 + 2 that are arched; an articular surface onmetacarpal I for metacarpal II that is nearly half the length ofthe former bone) but lacks others (the lateral shaft margin ofmetacarpal I is straight; metacarpal I is proportionally longfor its width; rectangular proximal end of metacarpal III),suggesting a primitive placement within, or along the stemto, Tetanurae. A second specimen from the same locality(SNGM-1888, 1889, 1895, 1901) includes an ilium, tarsusand pes that show a similar mixture of tetanuran (reducedsupraacetabular crest; pubic peduncle length:width ratio of1.6) and primitive (iliac pubic peduncle not substantiallylarger than ischial peduncle; oblique dorsal margin of astra-galar ascending process; low ascending process; proximalmetatarsal III lacking lateral notch) features.
Bittencourt & Kellner (2004) assigned an articulatedsequence of sacral and anterior caudal vertebrae from theSantana Formation of Brazil (MN 4743-V; Aptian–Albian)to Spinosauridae based on the absence of paired anteriorprocesses of the chevrons, an assignment with which weconcur. The caudal centra lack pleurocoelous fossae orforamina and show low but distinct neural arch laminaebelow the transverse processes. The neural spines are back-swept and anteroposteriorly narrow. In addition, a ‘supradi-apophyseal’ fossa is present near the base of the spine on thedorsal surface of the neural arch. Such fossae are known inthe caudals and posterior dorsals of Suchomimus and Bary-onyx but not in Torvosaurus or other non-spinosaurid mega-losauroids. The mediolaterally narrow centrum proportionsare also similar to those of Suchomimus but not Mega-losaurus, Torvosaurus or Eustreptospondylus.
The Late Cretaceous of South America has producednumerous, albeit often fragmentary remains of mega-losauroids in addition to the taxa described above.Spinosaurid teeth have been reported from the AlcantaraFormation of Maranhao (upper Cenomanian; Medeiros2005, 2006), the Cerro Lisandro Formation of RıoNegro (upper Cenomanian–lower Turonian; Canudo etal. 2004b), and the Adamantina Formation of Sao Paolo(Turonian–Santonian; Candeiro et al. 2004, 2006). Thelatter would be the youngest known record of the family.
Fragmentary carcharodontosaurid remains occur in theCerro Barcino Formation (Cerro Castano Member, Aptian)of Chubut (Vickers-Rich et al. 1999; possibly belong-ing to Tyrannotitan), as well as other sites in Argentina(Portezuelo Formation; these may pertain to Megarap-tor; Veralli & Calvo 2003, 2004). Purported carcharodon-tosaurid teeth from the Allen Formation of Argentina (?lateMaastrichtian; Martinelli & Forasiepi 2004) and the Marıliaand Adamantina Formations of Brazil (Late Cretaceous;Candeiro et al. 2004, 2006) exhibit enamel wrinkles thatare now known widely across Tetanurae (Brusatte et al.2007). However, the presence of apically inclined denticlesin these specimens, and strong similarity with the teeth of
the abelisaurid Skorpiovenator, suggest abelisaurid affini-ties (Canale et al. 2009).
Discussion
Comparisons with previous analysesAs is evident from the preceding sections, the number ofpublished analyses addressing the phylogenetic interrela-tionships of tetanuran theropods has become quite large(see online Supplementary File 2). It is neither practi-cal nor especially enlightening to review each individ-ually, given that even relatively minor changes in char-acter coding, taxon content, and search strategy canyield different topologies, each of which might poten-tially be scrutinized. Instead, we review several of themore contentious points of tetanuran phylogeny, as wellas certain common results, in the light of our presentanalysis.
Nonetheless, the underlying morphological observationsare the fundamental components of any analysis, and mustalso therefore be the most defensible. We have presentedour observations exhaustively in this work and its appen-dices, and encourage future workers to question any aspectof our basic data. Specific comparative comments withrespect to character identification, coding, and use canbe found among the descriptions provided in the OnlineSupplementary Material, along with a complete history ofeach character.
Carnosauria. A few recent studies have supported amonophyletic Carnosauria consisting of allosauroids,spinosauroids (megalosauroids) and an assortment ofclosely related taxa (Kurzanov 1989; Currie 1995; Rauhut2003), although only Rauhut (2003) provided explicit char-acter support. This arrangement effectively reconstitutesthe original concept of the group (Huene 1920) as a lineageof large bodied theropods excluding ceratosaurs and tyran-nosaurids (other studies have used the term Carnosauria asan approximate but not exact synonym of Allosauroidea,referring to a more restricted clade [Holtz et al. 2004]that is recovered in some form here). We did not recovera clade equivalent to Huene’s Carnosauria in this study,instead supporting monophyletic Allosauroidea and Mega-losauroidea as successive outgroups to Coelurosauria.
This is consistent with the results of a greater numberof studies (Holtz 1994a; Sereno et al. 1994, 1996; Sereno1999; Holtz et al. 2004) and is not surprising given therelatively low number of characters (3) previously foundto support Carnosauria unambiguously (Rauhut 2003). Allthree characters in the latter analysis (presence and dimen-sions of maxillary anterior ramus; opisthocoelous cervicalvertebrae; stout metacarpal I) are included here, but withaltered definitions and codings. Combined with improved
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taxon sampling, their support is no longer confined tothis grouping of taxa. (Our matrix requires at least sevenadditional steps to recover a monophyletic Carnosauria,and we find only two characters that provide unambigu-ous support for such a node: 45(2) and 177(2), the latterappearing in parallel in some ceratosaurs. A few other char-acters provide ambiguous support (131(1), 192(1), 226(1),237(1)). However, if one or more fragmentary forms ofuncertain placement (see branch support and taxon instabil-ity, above) are resolved with significantly different phyloge-netic positions than those proposed here, additional supportmight be garnered for a monophyletic Carnosauria.
Megalosauroidea. Unlike most previous analysis, werecovered a diverse and well supported Megalosauroidea,composed of three main clades. More commonly, mega-losauroids have been found to be a paraphyletic grade(Holtz 1994a; Charig & Milner 1997) or a clade consist-ing of relatively few taxa (Sereno et al. 1994, 1996, 1998;Sereno 1999). Concerted efforts to redescribe and restudypotential megalosauroids (Allain & Chure 2002; Rauhut2004; Sadleir et al. 2008; Benson 2008a, 2009a,b, 2010a,b),combined with important new discoveries (Sereno et al.1994; Allain 2002, 2005a; Rauhut 2005a), has allowed thediscovery of new character states and the coding of previ-ously indeterminate characters. This, in turn, has partlyresolved two of the main problems associated with mega-losauroid taxa: their rarity and their incompleteness.
Therefore we can offer considerable new support fora monophyletic Megalosauroidea, and for its constituentclades. The recognition of a relatively high taxonomicdiversity within this clade is an important step in under-standing the pattern and pace of theropod evolution. Italso provides a robust morphological characterization fora diverse, long-lived theropod clade, which should aid infuture identification of potential megalosauroid remains.Importantly, both Megalosauroidea and Megalosauria arewell supported by character optimization (Online Supple-mentary Table 1).
Allosauroidea. We have also found strong additionalsupport for a monophyletic Allosauroidea, with 11 unam-biguous characters resolved at this node plus many moreambiguous ones (Online Supplementary Table 1). Althoughmetriacanthosaurids share many features (e.g. 18(0), 72(0),107(0), 126(0), 136(0), 137(0), 138(0), 196(0), 219(0),224(1), 226(0), 272(1), 288(0), 297(0), 301(1), 320(0),324(1) with more basal tetanuran clades, their plesiomor-phic nature does little to weaken overall support becausethere are few competing synapomorphies that would specif-ically place them elsewhere in the phylogeny. However,if additional studies support the placement of Metriacan-thosauridae within Allosauroidea, this finding would indi-cate that many ‘advanced’ features of allosaurians werederived in parallel with Coelurosauria.
Support is also strong for the monophyly of the remain-ing allosauroids. This includes Allosauria, Carcharodon-tosauridae and Neovenatoridae. Carcharodontosauridae isstrongly supported, but within it we do not recover strongsupport for a sister-taxon relationship between Giganoto-saurus and Mapusaurus, which have been suggested toform the clade Giganotosaurinae (Coria & Currie 2006).At present we refer to the well-supported group includingthese two taxa and other derived carcharodontosaurids bythe name Carcharodontosaurinae, which has nomenclaturalpriority over other family-group names due to the inclusionof Carcharodontosaurus.
Taxa outside major clades. Our analysis revealsthat a number of basal tetanuran taxa exist outsidethe major clades (Megalosauroidea, Allosauroidea andCoelurosauria). We informally refer to these as ‘stem’ teta-nurans. Several of these taxa are relatively well represented,and are therefore unlikely to be excluded from major cladesdue to incomplete character data. As ‘stem’ taxa, theyprovide important temporal and morphological samples forunderstanding theropod evolution. ‘Stem’ taxa have beenuncommon finds in other analyses, which have producedresults that either favour multitaxon clades (Sereno et al.1994, 1996, 1998; Sereno 1999; Smith et al. 2007) or acomb-like topology overall (Holtz 1994a; Charig & Milner1997; Rauhut 2003; Holtz et al. 2004).
Two of these ‘stem’ taxa, ‘Dilophosaurus’ sinensis andCryolophosaurus, were previously found to form a cladeof their own along with Dilophosaurus wetherilli andDracovenator regenti (Smith et al. 2007), but this is notsupported in the present study. Our exclusion of severalbasal theropods, including Dracovenator and Zupaysaurus,may have contributed to this difference, but more likely itresults from distinctions in character choice and coding. Thethree synapomorphies of the ‘dilophosaur’ clade (Smith etal. 2007) are all related to the presence of a nasolacrimalskull crest that at least partly includes the premaxilla andinvolves pneumaticity of the lateral nasal. However, thelatter character is widely present among theropods anddiffers only in degree; we know of no theropods withany nasal crest development in which pneumaticity is notinvolved. It is not recovered as a unique synapomorphy ofthese taxa here.
Character polarity issues. Although previous studieshave noted numerous features shared between allosauroidsand coelurosaurs based on the osteology of Allosaurus (e.g.Paul 1984, 1988a, b), our study indicates that many ofthese were independently derived among allosaurians andbasal coelurosaurs. Thus, use of Allosaurus as an outgroupin studies of coelurosaur interrelationships may give anerroneous indication of character polarity by imperfectlyrecording the basal morphology for Allosauroidea (andAvetheropoda).
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Instead, we recommend that future studies ofCoelurosauria use relatively complete, basal representa-tives of Tetanurae, Megalosauroidea and Allosauroideaas outgroups. These include Sinraptor, Piatnitzkysaurus,Marshosaurus and Monolophosaurus, and possibly morebasal taxa such as Cryolophosaurus and ‘Dilophosaurus’sinensis. Several of these taxa have been described in detail(Currie & Zhao 1994; Smith et al. 2007; Brusatte et al.2010a; Zhao et al. 2010) but many have not; descriptionsof ‘D.’ sinensis, Piatnitzkysaurus and Marshosaurus remaina priority as these taxa may be exemplars of anatomicalevolution on the line leading to Coelurosauria (and Aves).More derived allosauroids and megalosauroids, includingwell-known taxa such as Allosaurus and Torvosaurus, likelypossess numerous derived features that obscure characteroptimizations for more inclusive clades. Finally, we wish tounderscore the importance of making direct observationson specimens, rather than relying primarily on publishedreports that will almost certainly never fully represent theavailable sample.
Evolutionary implicationsSkull design. Two basic skull morphologies are evidentwithin basal tetanurans. The first, usually considered typi-cal for large theropods, is exemplified by Allosaurusand Sinraptor. The skull is 2.5–3 times longer thantall. The snout tends to be fairly blunt and elabora-tions (e.g. horns and crests) are frequently present alongvarious parts of the skull roof, especially the lacrimal,nasals and frontals. This skull morphology is common inallosauroids, Cryolophosaurus and (outside of Tetanurae)within ceratosaurs. It may be primitive for tetanurans.
The second type, best observed in megalosauroids(e.g. Torvosaurus, Marshosaurus and Dubreuillosaurus),is characterized by a lower and longer skull, exhibitinga length:height ratio that can exceed 3:1. The skull roofis much less elaborated, with horns and crests being eithersmall or entirely absent, even in large individuals. The snoutregion in particular tends towards elongation although itremains blunt in some forms. Spinosaurids, although theyevolved a highly specialized skull morphology of their own,can be seen as having been derived from this basic morphol-ogy, and retain some of its main aspects of proportion anddesign.
The posterior skull is little modified in most tetanurans,especially when compared to the anterior and dorsal regionsof the skull. The primary exception is within Spinosauridae,in which the posterior skull has been elongated dorsoven-trally so as to extend below the ventral margin of themaxillary tooth row. Correspondingly, the basipterygoidprocesses are located ventral, rather than anterior, to thebasal tubera, suggesting that the posterior palate was alsolowered, resembling the condition in crocodiles. In addition,this region is bowed out laterally at its ventral edge, creat-ing a trapezoidal shape that contrasts with the more rectan-
gular cross section of other tetanuran skulls. Consideringthe remarkable degree of snout elongation in spinosaurids(Sereno et al. 1998; Dal Sasso et al. 2005), the braincase isextremely short anteroposteriorly.
Spinosaurids also exhibit the only significant departurefrom the ‘typical’ theropod tooth morphology among basaltetanurans, again converging on a crocodile-like form. Thepresence of paired tooth positions separated by diastem-ata also occurs uniquely within Spinosauridae amongtheropods. Elsewhere within Tetanurae, there are fewersignificant differences in tooth morphology or arrangement.Although subtle distinctions can be made between certaintetanuran groups (e.g. carcharodontosaurines), it is not clearwhether these indicate any substantial divergence in preda-tory or feeding habits.
Locomotor morphology. In some features, basal teta-nurans record transitions between the most primitivetheropod morphologies and the more derived, bird-likestates observed in coelurosaurs. For example, mega-losauroids, some allosauroids (including certain individualsof Allosaurus; Benson 2009b) and basal coelurosaurs suchas Tugulusaurus (Rauhut & Xu 2005) show an intermediatecondition in the orientation of the femoral head between 45◦
anteromedial (as in coelophysoids, ceratosaurs and moreprimitive forms) and fully medial (as in most avetheropods).Additional variation occurs in femoral head orientation;for example, the head is proximomedially inclined in mostcarcharodontosaurians, whereas in Neovenator it is orientedanteromedially at 45◦, similar to the condition in non-tetanurans (and still, also inclined proximally; Brusatteet al. 2008). The ‘proximal articular groove’, an obliquefurrow on the proximal surface of the femoral head, is alsopresent in megalosauroids and other basal tetanurans, incommon with non-tetanuran theropods. It is subsequentlylost and remains absent in avetheropods, including Neove-nator. Similarly, these taxa exhibit disparity in the size ofthe peduncles of the ilium, although less marked than incoelurosaurs. The latter occurs by reduction of the ischialpeduncle, accompanied by increase in relative anteroposte-rior length and absolute size of the pubic peduncle. Pedunclesize disparity is similar in megalosauroids and metriacan-thosaurids, suggesting that the coelurosaur-like features ofderived allosauroid peduncles (e.g. Aerosteon: Sereno et al.2008) appeared independently.
In other locomotory features, however, most basal teta-nurans had already achieved the derived condition. Theseinclude the presence of a laminar astragalar ascend-ing process, anteriorly directed distal astragalar condyles,some degree of reduction of the supraacetabular shelf(except in Chuandongocoelurus, Monolophosaurus andmore basal taxa) and absence of the shelf ‘morph’ of the M.iliofemoralis insertion on the proximal femur (‘trochantericshelf’; also still present in very basal tetanurans).
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Although variations occurred in the locomotor morphol-ogy of basal tetanurans, these appear to have been ofa much lesser degree relative to certain other theropodclades (e.g. ornithomimids, therizinosauroids, noasauridsand abelisaurids). Indeed, basal tetanurans exhibit a rela-tively generalized locomotor morphology among theropodsoverall. Specifically, size independent tendencies towardseither cursoriality or graviportality in limb proportions(Carrano 1999) are rare among most allosauroids andmegalosauroids. The single exception is among megara-ptorans, which evolved a proximally placed femoral lessertrochanter, elongate distal hind limb elements and a tallerascending process of the astragalus (Benson et al. 2010),all suggestive of cursoriality, although whether related tospeed or vagility is not determinable (Carrano 1999). Modi-fications of the pes comparable to those seen in somecoelurosaurs are likewise absent among more basal teta-nurans (Holtz 1994b).
Body size. The evolution of body size in dinosaurs hasbeen discussed informally for decades but only approachedanalytically in recent years (e.g. Hone et al. 2005; Carrano2006). Although approaches have differed, recent work-ers have agreed generally on the consistent pattern ofbody size increases within most groups of dinosaurs, withnotable exceptions being coelurosaurian theropods andtitanosaurian sauropods (e.g. Carrano 2006).
More broadly within theropods, however, patterns aremore complex. The most primitive theropod, Eoraptor, wassmall (tens of kilograms), but herrerasaurids were morethan five times larger than this. Basal coelophysoids (suchas Dilophosaurus and Zupaysaurus) were similar in sizeor even larger than herrerasaurids, indicating that morederived coelophysids underwent a size decrease. A dualpattern is evident within ceratosaurs, with large basal forms(Elaphrosaurus and Ceratosaurus, both in the 1
2 –1 tonnerange) yielding to descendants that were both considerablysmaller (noasaurids) and somewhat larger (abelisaurids)(Carrano & Sampson 2008).
Basal tetanurans were the first theropod clade to regu-larly achieve truly giant body sizes, with both mega-losauroid and allosauroid taxa weighing more than 1tonne. Even most smaller basal tetanuran taxa werefairly large, approximately the size of Ceratosaurus orDilophosaurus. Although Chuandongocoelurus, estimatedat 13 kg (Brusatte et al. 2010c), is an exception, it isunclear whether the holotype represents an adult indi-vidual. Interestingly, although both megalosauroids andallosauroids show evidence of size increases, neither cladeexhibits any evidence of marked size decreases (i.e. morethan an order of magnitude). Furthermore, within eachof these clades, there are multiple instances of majorsize increases. Giant (≥1 tonne) taxa include members ofboth Spinosauridae (Suchomimus, Spinosaurus) and Mega-losauridae (Torvosaurus) within Megalosauroidea, as well
as Allosauridae (Saurophaganax), Carcharodontosauridae(e.g. Carcharodontosaurus, Giganotosaurus, Mapusaurus,Tyrannotitan) and Neovenatoridae (Chilantaisaurus) withinAllosauroidea.
The apparently sequential temporal appearances of suchincreases led Bakker et al. (1992) to suggest that they repre-sented a successive series of ‘size-cycles’ of large theropods(femur length > 1100 mm): yangchuanosaurs (= metri-acanthosaurids), megalosaurs, allosaurs, acrocanthosaursand tyrannosaurs. They noted that although each replac-ing theropod group was more bird-like than the ‘incum-bent’ family, replacement depended on the extinction of theincumbent giant forms. The giant North African theropods(Spinosaurus, Bahariasaurus and Carcharodontosaurus)were considered to be a parallel radiation of aquatic preda-tors.
This size-cycle hypothesis is significantly complicated bynew discoveries and new phylogenetic hypotheses as wellas refined dating of theropod bearing horizons. The earliestlarge (> 1 tonne) theropods are still the early Late Jurassicmetriacanthosaurids, but the largest allosaurids are approx-imately coeval with giant megalosaurs in the upper levels ofthe Morrison Formation (Foster 2003), as well as the large(unidentified) theropods of the Tendaguru. Giant carchar-odontosaurians (including ‘acrocanthosaurs’) occur fromAptian through at least Coniacian times, contemporaries ofthe giant terrestrial spinosaurids. The largest tyrannosauridsdo not occur until the Campanian–Maastrichtian, when thelargest abelisaurids are also known from South America.Thus it was quite possible for more than one giant thero-pod to coexist, even in the same palaeoenvironment, butthis may have been dependent on significant variance infeeding habit between taxa.
Finally, although understanding the evolution of largesize has greater scientific value than the absolute sizes oftheropods per se, it is the latter that has garnered almostthe entirety of popular and scientific attention since theoriginal discovery of Tyrannosaurus rex. The emphasis onabsolute sizes has had the unfortunate effect of sideliningmany questions of genuine biological interest, including thepattern and process of size increases, their overall magni-tude (i.e. from ancestor to descendant), apparent repetition,and possible upper limits.
As an exception, Bakker et al. (1992) focused on howeach large theropod family independently reached nearlythe same maximum size. This remains true even in light ofthe many new discoveries of giant theropod remains madesince. Indeed, the difference in size between the skulls of thelargest carcharodontosaurids and tyrannosaurids amountsto only a few centimetres, well within the range of speciesvariation and rendering scientifically moot the question ofwhich is the ‘largest theropod’. Interestingly, these taxaexhibited different overall body dimensions, fore- andhind limb morphologies, and tooth shape, which impliesquite different predatory habits. Body mass (rather than
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Acrocanthosaurus
SaurophaganaxAllosaurus
Poekilopleuron
Eustreptospondylus
Piveteausaurus
Metriacanthosaurus
Condorraptor
Afrovenator
‘D.’ sinensisCryolophosaurus
BaryonyxSuchomimus
Spinosaurus
CarcharodontosaurusTyrannotitan
Angaturama
Neovenator
S. hepingensis
GiganotosaurusMapusaurus
Y. zigongensis
Megaraptor
Siamotyrannus
Hidden Lake
Fukuiraptor
S.dongi
Chuandongocoelurus
Aerosteon
Australovenator
Chilantaisaurus
Eocarcharia
Megalosaurus
Marshosaurus
Erectopus
Duriavenator
Monolophosaurus
Piatnitzkysaurus
TorvosaurusLeshansaurus
Dubreuillosaurus
IrritatorXuanhanosaurusShidaisaurus
Y. shangyouensis
Shaochilong
COELUROSAURIA
sampling
Streptospondylus
Concavenator
1
2
3
4
5
6
7
8
9
Magnosaurus
Orkoraptor
Jurassic CretaceousEarly Middle Late Early Late
He Si Pl To Aa Bj Bt Cl Ox Ki Ti Be Va Ha Ba Ap Al Ce Tu Co Sa Ca Ma
Figure 13. Stratigraphically calibrated phylogeny, based on current results. Note that coelurosaurs and taxa outside Tetanurae have beenomitted for clarity. Solid bars represent range estimates (including uncertainty) for taxa included in the present study; open bars representrange extensions based on taxa and specimens assigned to the respective clades; solid lines represent missing lineages inferred from thetopology; dashed lines represent tentative assignments. Filled circles along time line represent known theropod temporal occurrences, basedon 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 spec-imens exhibiting limb proportions that suggest substantiallyheavier masses than carcharodontosaurids of similar lineardimension. Bipedalism may have constrained the maximumsize attainable by theropods (Bakker et al. 1992; Farlowet al. 1995), but this hypothesis requires further testing.
Diversity. At a minimum, tetanurans originated early in theEarly Jurassic (Fig. 13), as evidenced by ‘Dilophosaurus’sinensis and Cryolophosaurus, as well as the sister taxonof Tetanurae, Ceratosauria, which is first representedby Berberosaurus (Pliensbachian–Toarcian; Allain et al.2007). Given the presence of coelophysoids in the LateTriassic, this implies the existence of stem-averostrans inthe Carnian. Of course, as this is a minimum divergenceestimate 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 themethod of Marshall 1994, which does not assume randomdistribution of samples) (Fig. 14).
Regardless of the exact time of origin, the earliest teta-nurans diversified alongside two other distinct theropodclades, coelophysoids and ceratosaurs. These three cladescoexisted through the Early Jurassic, but coelophysoidsapparently went extinct at the close of this interval or shortlythereafter (Carrano & Sampson 2004). Neither ceratosaursnor tetanurans appear to have been particularly diverseduring the Early Jurassic, but the poor quality of the fossilrecord from this interval, especially in terms of geograph-ical coverage, suggests that this may be partly artefactual.
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The phylogeny of Tetanurae 271
Jurassic CretaceousEarly Middle Late Early Late
He Si Pl To Aa Bj Bt Cl Ox Ki Ti Be Va Ha Ba Ap Al Ce Tu Co Sa Ca Ma
‘D.’ sinensisCryolophosaurus
Hidden Lake
Chuandongocoelurus
Erectopus
Monolophosaurus
COELUROSAURIA
Allosauridae
Metriacanthosauridae
Neovenatoridae
Spinosauridae
Carcharodontosauridae
Piatnitzkysauridae
Megalosauridae
Figure 14. Stratigraphically calibrated phylogeny showing sampling density for major clades, simplified from Fig. 13. Black circlesrepresent individual occurrences; white circles represent tentative assignments; black bars represent densely sampled intervals; white barsextending beyond noted samples represent range uncertainties.
The minimum divergence times of subsequent tetanuranclades suggest a radiation no later than the late Early Juras-sic (Fig. 13).
Subsequent to their initial appearance, Tetanurae radiatedinto two main clades, Megalosauroidea and Avetheropoda.The earliest megalosauroids are known from the Bajocian(Duriavenator, Magnosaurus), implying the presence ofbasal avetheropods at this time also. Definitive avethero-pod fossils occur slightly later in the Middle Jurassicof Europe (Bathonian: Proceratosaurus). In the MiddleJurassic of China, ‘stem’ tetanurans (Chuandongocoelu-rus, Monolophosaurus) persisted alongside megalosaurids(Leshansaurus) and metriacanthosaurid allosauroids, thefirst occurrences of which (Shidaisaurus) are penecontem-poraneous with those of avetheropods in Europe.
By the Callovian and Late Jurassic, the fossil recorddemonstrates the widespread presence of multiple cladeswithin both megalosauroids and avetheropods. Amongthe former are megalosaurids and piatnitzkysaurids,implying the presence of spinosaurids as well. Indeed,Eustreptospondylus and Streptospondylus are ambiguouslyresolved as basal megalosaurians and may therefore repre-sent the most basal spinosaurids (although they couldequally well be megalosaurids). Avetheropodans includedallosaurids and metriacanthosaurids, therefore implying thepresence of basal carcharodontosaurians, which may berepresented in the Tendaguru Formation of Tanzania. Theearliest definite spinosaurid fossils are possibly Late Juras-sic (Tendaguru Formation) but certainly mid-Neocomianin age; the earliest definite carcharodontosaurian fossils
are possibly also Late Jurassic (Tendaguru Formation), butcertainly Barremian (Concavenator, Neovenator). Fossilsof both groups are abundant in the ‘medial’ Cretaceousinterval.
Allosaurids are an unexpectedly small group, restricted toAllosaurus and Saurophaganax and present with certaintyonly in the Kimmeridgian–Tithonian of North America andEurope. Metriacanthosaurids are almost exclusively knownfrom the Oxfordian of Asia and Europe. It is not yet possibleto determine whether metriacanthosaurids were present inthe Oxfordian of North America but replaced by allosauridsin the Kimmeridgian, as seems to have occurred in Europe.Alternatively, the distributions of these two clades mayreflect genuine biogeographical differentiation.
Among the megalosauroids, only spinosaurids defini-tively survived the Early Cretaceous, although there mayhave been a relictual megalosauroid in Late CretaceousAntarctica (Hidden Lake specimen). Afrovenator was thesole non-spinosaurid megalosauroid known for this inter-val, but recent work suggests that it may be considerablyolder (Rauhut & Lopez-Arbarello 2009). Although theirinitial diversification included nearly all the continents,megalosauroids may have been restricted to Europe andGondwana by the late Early Cretaceous. The paucity ofspinosaurid discoveries from the late Early and early LateCretaceous may reflect genuinely sparse and geographicallypatchy fossil sampling. Allosauroids were also geograph-ically widespread in their early history, and they tooare primarily known from Gondwanan landmasses bythe 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 atca 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 andquestionable or indeterminate basal tetanurans; squares, megalosauroids; triangles, allosauroids. Maps copyright Ron Blakey, NorthernArizona University.
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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; greycircles, megalosauroids; filled circles, avetheropods.
are known from the ?Turonian of China, but the recordof pre-Campanian Late Cretaceous theropods from Laura-sia is otherwise very poor. It is likely that the Laurasianradiation of large bodied coelurosaurs, known primar-ily from the Campanian and Maastrichtian, came atthe eventual expense of more basal tetanuran lineages.South American megaraptoran allosauroids are the onlyCampanian–Maastrichtian basal tetanurans.
Finally, it is interesting to note how common it isto find multiple lineages of apparently contemporane-ous large theropods within single formations of Juras-sic and Early Cretaceous age. Although the MorrisonFormation of western North America has been noted forits unusually high diversity of large theropods (at leastfive; Chure et al. 2000), other locales have producedat least three contemporaneous taxa. In addition tothe Late Jurassic beds of Tanzania and Portugal, theseinclude the Bathonian (a megalosaurid and basal tetanu-rans) and Berriasian (allosauroids, a spinosaurid) of theUnited Kingdom, the Albian and Cenomanian of NorthAfrica (spinosaurids, carcharodontosaurids, ceratosaurs),the Allen Formation of Argentina (basal tetanurans,carcharodontosaurids, abelisaurids), and perhaps theAdamantina Formation of Brazil (?spinosaurids, ?carchar-odontosaurids, abelisaurids). Many additional formationsdocument the presence of at least two large theropods, butmost of these are far from exhaustively sampled.
Biogeography. The biogeographical history of varioustheropod groups has received intense attention in recentyears (e.g. Carrano et al. 2002; Sereno et al. 2004), butlittle consensus has been achieved. It is beyond the scopeof this paper to review the major hypotheses and anal-yses, even briefly. Unlike phylogenetic analyses, there aresignificant discrepancies in terminology, data sampling, and
methodology that make comparisons between results prob-lematic. Instead, we use the results of the present phylo-genetic analysis to suggest a substantial revision to thenull models currently employed by many biogeographicalstudies.
As noted in a previous paper (Benson et al. 2010), mosttheropod clades achieved a broad geographical distributionquite early in their geological history. It is this early historythat should inform considerations of what the neutral expec-tations should be for any biogeographical scenario. Withthis in mind, we suggest that any diverse clade appearingprior to the Late Jurassic should be suspected of havinga global distribution a priori, at least on the continen-tal geographical scale. Given the incompleteness of thefossil record, it is therefore especially important to deter-mine genuine absences, because well-supported evidenceof absences will be particularly enlightening. For exam-ple, the absences of sauropods and large non-tyrannosauridtheropods from the Hell Creek Formation of North Amer-ica appear to be genuine, based as they are on the lack offossils of these forms among the hundreds of thousandscollected thus far. The presence of both groups in earlierstrata, and their survival elsewhere in the Maastrichtian,indicates that a regional extinction is responsible for thispattern.
Regional extinctions must be more important than hasbeen suspected in determining the later Mesozoic patternsof dinosaur distribution. As discovery and analysis haveadvanced in recent decades, most known clades have beenidentified in progressively older strata, and indeed nearly allmajor dinosaur clades (e.g. Ceratopsia) are now known frompre-Late Jurassic rocks. Theropods conform to this trend,with definite coelurosaurs (and therefore all more primi-tive theropod clades) now well identified from the MiddleJurassic, and several clades that probably originated in theJurassic (metriacanthosaurids, megalosauroids, carcharo-dontosaurians) showing an early widespread distribution(Fig. 15).
In this respect, the modern world and the distributionsof its terrestrial taxa constitute a poor model for Meso-zoic continental biogeography, because most extant cladeshad their origins in a world of fragmented continents.Even so, many modern clades whose origins are knownto predate significant continental breakup (e.g. Marsu-pialia/Metatheria, Pleurodira, Anguimorpha) have fossilrecords that are nearly global in breadth.
We suggest a model wherein the initial supposition is forlarge Mesozoic clades to have had the potential to achievea cosmopolitan distribution, which would befit an earlyMesozoic world in which continental connection was therule. This operates under the assumption that organismsquickly reach their maximum potential dispersion, at leaston geological time frames, and sets the groundwork forinterpreting observed taxon distributions. It does not imply
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The phylogeny of Tetanurae 275
that all dinosaur clades had global distributions; rather it isa first-order expectation that provides a framework againstwhich deviations can be identified and studied.
Second, it is critical to assess the quality of thefossil record. Given recent advances in understanding thedinosaur fossil record and the steady rate of discovery ofnew taxa, it is not tenable merely to assume that all taxonabsences are real in every locale. The tendency to do soinvolves the tacit assumption that the fossil record is mostlycomplete and/or accurate, and therefore does not require asrigorous analytical treatment as phylogenetic hypotheses.(One would not support coding every ‘?’ as ‘0’ in a phylo-genetic analysis, but this is analogous to assuming all taxonabsences are genuine in the fossil record.)
With this information in hand, it becomes possible toexamine biogeographical patterns. Deviations from priorexpectations can be analysed in more detail in the light ofthe known fossil record. Only at this stage is it appropriateto introduce second-order geographical barriers (e.g. moun-tain ranges, deserts, or other features that impose differen-tial restrictions on different terrestrial organisms). We knownext to nothing about the actual dispersal potentials for anydinosaurs with respect to barriers of this kind, and so theycannot be invoked as independent causative factors unless itcan first be demonstrated that taxon absences are likely to bereal. This is not to say that such barriers were not importantin creating biogeographical patterns amongst dinosaurs inthe Mesozoic – certainly they were. But our knowledge ofthem, and of how dinosaurs interacted with them, is insuf-ficient to permit them a role in setting prior expectations.
Stratigraphical fit. As with other dinosaur clades, thereis relatively good overall fit between the sequence ofclade appearances and their stratigraphical first appearanceamong tetanurans. This is reflected in the highly significantcorrelation of age rank and clade rank based on our phylo-genetic hypothesis (rho = 0.73, p < 0.001), as well as thegraphical comparison of these two variables (Fig. 16) andthe SCI of 0.605. The RCI is quite low (70.77%), emphasiz-ing how much of the predicted record of tetanurans remainsto be discovered.
Few major discrepancies occur in sequence; rather thereare extensive missing lineages throughout the phylogeny.Significant among these are the lineages leading toSpinosauridae (∼ 42 Ma missing), Allosauria (∼ 20 Ma),Spinosaurinae (∼ 20 Ma), and Carcharodontosauria (∼ 23Ma). The sampling gaps in late Early–early Middle andearliest Cretaceous times are apparent in the absence oftaxa with appropriate age ranks (7–8 and 15–16, respec-tively). The late appearing taxa with low clade rank arederived ceratosaurs, whose ranks are artificially low due tothe low numbers of taxa sampled for this study.
In addition, the two major ‘pulses’ of tetanuran evolu-tion (whether genuine or artefactual) are evident as discreteclusters of taxa in the centre and upper-right portions of
the graph. These reflect the Jurassic radiation of mega-losauroids and the Cretaceous radiations of spinosauridsand derived allosaurians.
Conclusions
Tetanuran theropods represent the majority of predatorydinosaurs, but their early evolutionary history has remaineddifficult to decipher. Although many taxa are known fromfragmentary remains, several recent discoveries combinedwith a re-examination of existing materials has allowedus to significantly improve phylogenetic resolution forthis important group. Here we present the results of adetailed, species-level phylogenetic analysis of 59 ingrouptaxa, mostly tetanurans. Our study utilizes 351 charactersderived from firsthand study of specimens and comprehen-sive review of all prior systematic analyses.
The results support the successive placement ofCeratosauria and Tetanurae as more derived groups rela-tive to Coelophysoidea. Several taxa are found to occupy‘stem’ positions relative to Tetanurae, including two(Cryolophosaurus and ‘Dilophosaurus’ sinensis) that werepreviously considered to be more basal. Within Tetanu-rae, Megalosauroidea and Avetheropoda are sister taxa.The former clade is quite diverse, with two Jurassicclades (Piatnitzkysauridae fam. nov. and Megalosauridae)and the Cretaceous Spinosauridae. Within Megalosauri-dae, our results support the presence of two subfami-lies, Megalosaurinae and Afrovenatorinae subfam. nov.Avetheropoda includes the Jurassic Metriacanthosauridaeand Allosauridae as well as the Cretaceous Carcharo-dontosauria (comprising Neovenatoridae and Carcharodon-tosauridae).
The presence of Early and Middle Jurassic forms onthe tetanuran ‘stem’ lineage requires a minimum appear-ance time of Late Triassic or earliest Jurassic for Averostra,and the early Middle Jurassic appearance of several truetetanurans (Duriavenator, Magnosaurus) suggests that theradiation of basal megalosauroids and avetheropods tookplace not later than the late Early Jurassic. Given the rela-tively poor Early Jurassic record, it is certainly possible thatthese lineages extend considerably farther back in time.Substantial missing lineages are present for Spinosauridaeand Allosauria, although the > 20 million-year gap forCarcharodontosauria may be mitigated by the presence ofa Tendaguru form.
In addition, we review all previously published tetanu-ran taxa and occurrences in order to document thoroughlyand assess the fossil record of the group. Many of thesetaxa were too fragmentary to be included in our analysis,but can nonetheless be identified as belonging to Tetanuraeor particular ingroups. Although some records of tetanu-rans can be refuted, the remaining sample documents amuch broader geographical and temporal radiation than the
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phylogenetic results alone. A revised classification of teta-nurans is given in the Appendix.
Tetanuran evolution seems to exhibit a series of ‘waves’of diversification, but it is difficult to determine whetherthese are relatively discrete pulses or an artefact ofuneven sampling. Regardless, the dominance of particulargroups, particularly at large body sizes, follows a loosesuccession: Megalosauridae, Piatnitzkysauridae andMetriacanthosauridae (Middle Jurassic); Allosauridae,Megalosauridae and Metriacanthosauridae (Late Jurassic);Spinosauridae, Carcharodontosauridae and Neovenatori-dae (Early and mid Cretaceous). Terminal Cretaceousterrestrial ecosystems appear to have been dominatedby a combination of large-bodied tetanurans (i.e. tyran-nosaurid coelurosaurians; primarily Northern Hemisphere)and abelisaurid ceratosaurians (primarily SouthernHemisphere).
The morphological evolution of tetanurans is complex, aswould be expected of a radiation involving several contem-poraneous clades. General skull proportions vary betweena more primitive design with greater skull roof elabora-tion (common in allosauroids and outside Tetanurae) anda more elongate form that lacks these developments (moretypical among megalosauroids). Spinosaurids show the onlysignificant departures in tooth morphology, snout configu-ration, and morphology of the posterior skull within non-coelurosaurian tetanurans. Although some basal tetanurans(e.g. Neovenatoridae) display parallels with coelurosaurs,in general the group is characterized by a relatively conser-vative limb morphology that shows little tendency towardsthe extremes of locomotor evolution. Tetanuran evolutionrecords a gradual transition from a primitive theropodcondition to the more bird-like features common amongcoelurosaurs, although a number of ‘avian’ features werealready present in many tetanuran taxa. Basal tetanuransachieved giant body sizes in a number of lineages, reach-ing very similar maximum sizes despite this independentevolution. The presence of multiple large theropods inseveral different palaeoenvironments suggests some degreeof ecological resource partitioning.
The biogeographical history of tetanurans is likewisecomplex, spanning more than 110 Ma and all continents.At present the density of sampling is insufficient, bothtemporally and geographically, to permit detailed anal-ysis of tetanuran biogeographic evolution. However, wesuggest an alternative model for assessing the history ofthe group, and of terrestrial vertebrates more generallyduring this time interval. The presence of all major lineagesof tetanurans prior to any significant breakup of Pangaeaimplies that these clades would have had the opportunityto disperse widely and achieve cosmopolitan distributions.Their absences from regions later in time would then bedue to regional extinctions or dispersal failure. This modelshould be appropriate for any early Mesozoic radiation;
a dispersal/vicariance dichotomy is an appropriate ‘nullmodel’ only for more highly nested groups whose originspostdate continental fragmentation.
Acknowledgements
We would like to thank the many people who allowedus access to specimens in their care, including: PeterWellnhofer (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 Xijinand Xu Xing (IVPP); John Foster (MWC); Jose Bona-parte, Alejandro Kramarz and Fernando Novas (MACN);Rodolfo Coria (MCF); Leonardo Salgado (MC); DiegoPol (MCF); Sean Duran (Miami Science Museum); SteveHutt and Martin Munt (MIWG); Octavio Mateus (ML);Philippe Taquet, Daniel Goujet, Ronan Allain and EmilyLong (MNHN); Rodolfo Coria (MPEF); Jorge Calvo(MUCPv); Juan Canale (MUCPv-Ch); Vince Schneiderand Drew Eddy (NCSM); Jeff Person (OMNH); PaulJeffery (OUMNH); Jaime Powell (PVL); Oscar Alcoberand Ricardo Martınez (PVSJ); Jaime Powell (PVL); DavidMullin (SDM); Rich Ketcham and Tim Rowe (TMM);Sandra Chapman, Angela Milner and Paul Barrett (NHM);Robert Masek and Paul Sereno (University of Chicago); PatHolroyd and Kevin Padian (UCMP); Mike Getty (UMNH);and Guang-Zhou Peng (ZDM). In several cases, we weregranted access to then-unpublished materials, for which weare particularly grateful. This paper benefited from numer-ous discussions with Jeffrey Wilson, Cathy Forster, RonTykoski, Paul Barrett, David Norman, and Nathan Smith,careful editorial work by Susannah Maidment, and detailedand insightful reviews by Oliver Rauhut and StephenBrusatte.
Translations of Allain (2001), Bonaparte & Novas(1985), Bonaparte (1986), Deperet & 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) andThevenin (1907) are available at the Polyglot Paleontol-ogist website (http://www.paleoglot.org/).
This research was supported by NSF DEB-9904045 toMTC and SDS; NERC studentship NER/S/A/2005/13488to RBJB; and smaller grants from The Jurassic Foundation,SYNTHESYS and the Palaeontographical Society to RBJB.
Supplementary material
Supplementary material is available online DOI:10.1080/14772019.2011.630927
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References
Aberhan, M., Bussert, R., Heinrich, W.-D., Schrank, E.,Schultka, S., Sames, B., Kriwet, J. & Kapilima, S.2002. Palaeoecology and depositional environments of theTendaguru Beds (Late Jurassic to Early Cretaceous, Tanza-nia). Mitteilungen aus dem Museum fur Naturkunde in Berlin,Geowissenschaftliche Reihe, 5, 19–44.
Adams, E. N., III. 1972. Consensus techniques and the compari-son of taxonomic trees. Systematic Zoology, 21, 390–397.
Agnolin, F. L., Ezcurra, N. D., Pais, D. F. & Salisbury, S. W.2010. A reappraisal of the Cretaceous non-avian dinosaurfaunas from Australia and New Zealand: evidence for theirGondwanan affinities. Journal of Systematic Palaeontology,8, 257–300.
Allain, R. 2001. Redescription de Streptospondylus altdorfen-sis, le dinosaure theropode de Cuvier, du Jurassique deNormandie. Geodiversitas, 23, 349–367.
Allain, R. 2002. Discovery of megalosaur (Dinosauria,Theropoda) in the Middle Bathonian of Normandy (France)and its implications for the phylogeny of basal Tetanurae.Journal of Vertebrate Paleontology, 22, 548–563.
Allain, R. 2005a. The postcranial anatomy of the megalosaurDubreuillosaurus valesdunensis (Dinosauria Theropoda)from the Middle Jurassic of Normandy, France. Journal ofVertebrate Paleontology, 25, 850–858.
Allain, R. 2005b. The enigmatic theropod dinosaur Erectopussuperbus (Sauvage 1882) from the Lower Albian of Louppy-le-Chateau (Meuse, France). Pp. 72–86 in K. Carpenter(ed.) The Carnivorous Dinosaurs. Indiana University Press,Bloomington.
Allain, R. & Chure, D. J. 2002. Poekilopleuron bucklandii, thetheropod dinosaur from the Middle Jurassic (Bathonian) ofNormandy. Palaeontology, 45, 1107–1121.
Allain, R., Tykoski, E., Aquesbi, N., Jalil, N.-E., Monbaron, M.,Russell, D. & Taquet, P. 2007. An abelisauroid (Dinosauria:Theropoda) from the Early Jurassic of the High Atlas moun-tains, Morocco, and the radiation of ceratosaurs. Journal ofVertebrate Paleontology, 27, 610–624.
Ambayrac, M. 1913a. Decouverte d’une machoire de reptileJurassique. Bulletin Mensuel des Naturalistes des Alpes-Maritimes, 15, 65–68.
Ambayrac, M. 1913b. Une machoire de grand reptile du Juras-sique superieur (Oxfordien). Association Francaise pourl’Avancement des Sciences, 42, 97–98.
Antunes, M. T. & Sigogneau-Russell, D. 1991. Nouvellesdonnees sur les dinosaures du Cretace superieur du Portu-gal. Comptes Rendus de l’Academie des Sciences a Paris,Serie II, 313, 113–119.
Antunes, M. T. & Sigogneau-Russell, D. 1992. La faunedes petits dinosaures du Cretace terminal portugais.Comunicacoes dos Servicos Geologicos de Portugal, 78,49–62.
Averianov, A. O., Martin, T. & Bakirov, A. A. 2005. Pterosaurand dinosaur remains from the Middle Jurassic BalabansaiSvita in the northern Fergana depression, Kyrgyzstan (centralAsia). Palaeontology, 48, 135–155.
Azuma, Y. & Currie, P. J. 2000. A new carnosaur (Dinosauria:Theropoda) from the Lower Cretaceous of Japan. CanadianJournal of Earth Sciences, 37, 1735–1753.
Bajpai, S. & Prasad, G. V. R. 2000. Cretaceous age for Ir-rich Deccan intertrappean deposits: palaeontological evidencefrom Anjar, western India. Journal of the Geological Society,London, 157, 257–260.
Bakker, R. T. 1986. The Dinosaur Heresies. William Morrow &Company, New York: 481 pp.
Bakker, R. T. 2000. Brontosaur killers: Late Jurassic allosauridsas sabre-tooth cat analogues. GAIA, 15, 145–158.
Bakker, R. T., Siegwarth, J., Kralis, D. & Filla, J. 1992.Edmarka rex, a new, gigantic theropod dinosaur from themiddle Morrison Formation, Late Jurassic of the Como Bluffoutcrop region. Hunteria, 2(9), 1–24.
Barriel, V. & Tassy, P. 1998. Rooting with multiple outgroups:consensus versus parsimony. Cladistics, 14, 193–200.
Barsbold, R. 1983. Khishchnye dinosavry mela Mongoliy.Sovmestnaya Sovetsko-Mongol’skaya PaleontologicheskayaEkspeditsiya, Trudy, 19, 1–117.
Baur, G. 1891. Remarks on the reptiles generally calledDinosauria. The American Naturalist, 25, 434–454.
Benson, R. B. J. 2008a. A redescription of ‘Megalosaurus’hesperis (Dinosauria, Theropoda) from the Inferior Oolite(Bajocian, Middle Jurassic) of Dorset, United Kingdom.Zootaxa, 1931, 57–67.
Benson, R. B. J. 2008b. New information on Stokesosaurus, atyrannosauroid (Dinosauria: Theropoda) from North Americaand the United Kingdom. Journal of Vertebrate Paleontology,28, 732–750.
Benson, R. B. J. 2009a. The Taxonomy, Systematics and Evolutionof the British Theropod Dinosaur Megalosaurus. UnpublishedPhD Thesis, University of Cambridge, 274 pp.
Benson, R. B. J. 2009b. An assessment of variability in dinosaurremains from the Bathonian (Middle Jurassic) of Stonesfieldand New Park Quarry, UK and taxonomic implications forMegalosaurus bucklandii and Iliosuchus incognitus. Palaeon-tology, 54, 857–877.
Benson, R. B. J. 2010a. A description of Megalosaurus bucklandii(Dinosauria: Theropoda) from the Bathonian of the UnitedKingdom and the relationships of Middle Jurassic theropods.Zoological Journal of the Linnean Society, 158, 882–935.
Benson, R. B. J. 2010b. The osteology of Magnosaurusnethercombensis (Dinosauria, Theropoda) from the Bajo-cian (Middle Jurassic) of the United Kingdom and a re-examination of the oldest records of tetanurans. Journal ofSystematic Palaeontology, 8, 131–146.
Benson, R. B. J. & Barrett, P. M. 2009. Dinosaurs of Dorset:Part I, the carnivorous dinosaurs (Saurischia, Theropoda).Proceedings of the Dorset Natural History and Archaeologi-cal Society, 130, 133–147.
Benson, R. B. J. & Radley, J. D. 2010. A new large-bodiedtheropod dinosaur from the Middle Jurassic of Warwickshire,United Kingdom. Acta Palaeontologica Polonica, 55, 35–42.
Benson, R. B. J. & Xu, X. 2008. The anatomy and systematic posi-tion of the theropod dinosaur Chilantaisaurus tashuikouensisHu, 1964 from the Early Cretaceous of Alanshan, People’sRepublic of China. Geological Magazine, 145, 778–789.
Benson, R. B. J., Barrett, P. M., Powell, H. P. & Norman, D.B. 2008. The taxonomic status of Megalosaurus bucklandii(Dinosauria, Theropoda) from the Middle Jurassic of Oxford-shire, UK. Palaeontology, 51, 419–424.
Benson, R. B. J., Brusatte, S. L., Hutt, S. & Naish, D. 2009. Anew large basal tetanuran (Dinosauria: Theropoda) from theWessex Formation (Barremian) of the Isle of Wight. Journalof Vertebrate Paleontology, 29, 612–615.
Benson, R. B. J., Carrano, M. T. & Brusatte, S. L. 2010.A new clade of archaic large-bodied predatory dinosaurs(Theropoda: Allosauroidea) that survived to the latest Meso-zoic. Naturwissenschaften, 97, 71–78.
Bibbins, A. 1895. Notes on the paleontology of the PotomacFormation. The Johns Hopkins University Circulars, 15,17–20.
Bidar, A., Demay, L. & Thomel, G. 1972. Sur la presence duDinosaurien Compsognathus dans le Portlandien de Canjuers
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
278 M. T. Carrano et al.
(Var). Comptes Rendus de l’Academie des Sciences, Paris,Serie D, 275, 2327–2329.
Bigot, A. 1945. La destruction des collections et des bibliothequesscientifiques de Caen. Bulletin de la Societe Linneenne deNormandie, Volume supplementaire, 1–75.
Bittencourt, J. S. & Kellner, A. W.A. 2004. On a sequenceof sacrocaudal theropod dinosaur vertebrae from the LowerCretaceous Santana Formation, northeastern Brazil. Arquivosdo Museu Nacional, Rio de Janeiro, 62, 309–320.
Bonaparte, C.-L. 1850. Conspectus Systematum Herpetologiae etAmphibiologiae. Editio Altera Reformata. E. J. Brill, Leyden.
Bonaparte, J. F. 1979. Dinosaurs: a Jurassic assemblage fromPatagonia. Science, 205, 1377–1378.
Bonaparte, J. F. 1986. Les dinosaures (Carnosaures,Allosaurides, Sauropodes, Cetosaurides) du JurassiqueMoyen de Cerro Condor (Chubut, Argentina). Annales dePaleontologie (Vert.-Invert.), 72, 247–289.
Bonaparte, J. F. 1991. The Gondwanian theropod familiesAbelisauridae and Noasauridae. Historical Biology, 5, 1–25.
Bonaparte, J. F. & Novas, F. E. 1985. Abelisaurus comahuensis,n.g., n.sp., Carnosauria del Cretacico Tardio de Patagonia.Ameghiniana, 21, 259–265.
Boneham, B. F. M. & Wyatt, R. J. 1993. The stratigraphical posi-tion of the Middle Jurassic (Bathonian) Stonesfield Slate ofStonesfield, Oxfordshire, UK. Proceedings of the Geologists’Association, 104, 123–136.
Bouaziz, S., Buffetaut, E., Ghanmi, M., Jaeger, J.-J., Martin,M., Mazin, J.-M. & Tong, H. 1988. Nouvelles decouvertesde vertebres fossiles dans l’Albien du sud tunisien. Bulletinde la Societe Geologique de France, 8e serie, 4, 335–339.
Bridge, J. S., Jalfin, G. A. & Georgieff, S. M. 2000. Geome-try, lithofacies, and spatial distribution of Cretaceous fluvialsandstone bodies, San Jorge Basin, Argentina: outcrop analogfor the hydrocarbon-bearing Chubut Group. Journal of Sedi-mentary Research, 70, 341–359.
Bristowe, A. & Raath, M. A. 2004. A juvenile coelophysoidskull from the Early Jurassic of Zimbabwe, and the synonymyof Coelophysis and Syntarsus. Palaeontologia Africana, 40,31–41.
Britt, B. B. 1991. Theropods of Dry Mesa Quarry (MorrisonFormation, Late Jurassic), Colorado, with emphasis on theosteology of Torvosaurus tanneri. BYU Geology Studies, 37,1–72.
Brokenshire, A. J. & Clarke, J. B. 1993. Important recentlycollected dinosaurian remains from the lower KimmeridgeClay at Weymouth. Proceedings of the Dorset Natural Historyand Archaeological Society, 115, 177–178.
Brusatte, S. L. & Sereno, P. C. 2007. A new species of Carchar-odontosaurus (Dinosauria: Theropoda) from the Cenomanianof Niger and a revision of the genus. Journal of VertebratePaleontology, 27, 902–916.
Brusatte, S. L. & Sereno, P. C. 2008. Phylogeny of Allosauroidea.Journal of Systematic Palaeontology, 6, 155–182.
Brusatte, S., Benson, R. B. J., Carr, T. D., Williamson, T. E. &Sereno, P. C. 2007. The systematic utility of theropod enamelwrinkles. Journal of Vertebrate Paleontology, 27, 1052–1056.
Brusatte, S., Benson, R. B. J. & Hutt, S. 2008. The osteol-ogy of Neovenator salerii (Dinosauria: Theropoda) from theWealden Group (Barremian) of the Isle of Wight. Palaeonto-graphical Society Monographs, 162, 1–75.
Brusatte, S., Benson, R. B. J., Chure, D. J., Xu, X., Sulli-van, C. & Hone, D. W. E. 2009a. The first definitivecarcharodontosaurid (Dinosauria: Allosauroidea) from Asia,the evolution of Asian Cretaceous theropods and the delayedascent of tyrannosauroids. Naturwissenschaften, 96, 1051–1058.
Brusatte, S. L., Carr, T. D., Erickson, G. M., Bever, G. S.& Norell, M. A. 2009b. A long-snouted, multihorned tyran-nosaurid from the Late Cretaceous of Mongolia. Proceed-ings of the National Academy of Sciences, 106, 17261–17266.
Brusatte, S., Benson, R. B. J., Zhao, X.-J. & Currie, P. J.2010a. The skull of Monolophosaurus jiangi (Dinosauria:Theropoda) and its implications for early theropod phylogenyand evolution. Zoological Journal of the Linnean Society, 158,573–607.
Brusatte, S., Chure, D. J., Benson, R. B. J. & Xu, X. 2010b. Theosteology of Shaochilong maortuensis, a carcharodontosaurid(Dinosauria: Theropoda) from the Late Cretaceous of Asia.Zootaxa, 2334, 1–46.
Brusatte, S., Benson, R. B. J. & Xu, X. 2010c. The evolution oflarge-bodied theropod dinosaurs during the Mesozoic in Asia.Journal of Iberian Geology, 36, 275–296.
Brusatte, S., Benson, R. B. J. & Xu, X. 2011. A reassessment ofKelmayisaurus petrolicus, a large theropod dinosaur from theEarly Cretaceous of China. Acta Palaeontologica Polonica(doi:10.4202/app.2010.0125).
Buckland, W. 1824. Notice on the Megalosaurus or great fossillizard of Stonesfield. Transactions of the Geological Societyof London, Series 2, 1, 390–396.
Buffetaut, E. 1982. Aggiosaurus nicaeensis Ambayrac, 1913,from the Upper Jurassic of south-eastern France: a marinecrocodilian, not a dinosaur. Neues Jahrbuch fur Geologie undPalaontologie Monatshefte, 1982, 469–475.
Buffetaut, E. 1989. New remains of the enigmatic dinosaurSpinosaurus from the Cretaceous of Morocco and the affini-ties between Spinosaurus and Baryonyx. Neues Jahrbuch furGeologie und Palaontologie Monatshefte, 1989, 79–87.
Buffetaut, E. 1992. Remarks on the Cretaceous theropoddinosaurs Spinosaurus and Baryonyx. Neues Jahrbuch furGeologie und Palaontologie Monatshefte, 1992, 88–96.
Buffetaut, E. 2007. The spinosaurid dinosaur Baryonyx(Saurischia, Theropoda) in the Early Cretaceous of Portugal.Geological Magazine, 144, 1021–1025.
Buffetaut, E. 2008. Spinosaurid teeth from the Late Jurassic ofTendaguru, Tanzania, with remarks on the evolutionary andbiogeographical history of the Spinosauridae. Pp. 26–28 inJ.-M. Mazin, J. Pouech, P. Hantzpergue & V. Lacombe (eds)Mid-Mesozoic Life and Environments. Cognac, France.
Buffetaut, E. 2010. Spinosaurs before Stromer: early finds ofspinosaurid dinosaurs and their interpretations. Pp. 175–188in R. Moody, E. Buffetaut, D. Naish & D.M. Martill (eds)Dinosaurs and Other Extinct Saurians: A Historical Perspec-tive. Geological Society Special Paper, 343, The GeologicalSociety, London.
Buffetaut, E. & Enos, J. 1992. Un nouveau fragmentcranien de dinosaure theropode du Jurassique des VachesNoires (Normandie, France): remarques sur la diversitedes theropodes jurassiques europeens. Comptes Rendus del’Academie des Sciences a Paris, Serie II, 314, 217–222.
Buffetaut, E. & Ingavat, R. 1986. Unusual theropod teeth fromthe Upper Jurassic of Phu Wiang, northeastern Thailand.Revue de Paleobiologie, 5, 217–220.
Buffetaut, E. & Ouaja, M. 2002. A new specimen of Spinosaurus(Dinosauria, Theropoda) from the Lower Cretaceous ofTunisia, with remarks on the evolutionary history of theSpinosauridae. Bulletin de la Societe Geologique de France,173, 415–421.
Buffetaut, E. & Suteethorn, V. 2007. A sinraptorid theropod(Dinosauria: Saurischia) from the Phu Kradung Formation ofnortheastern Thailand. Bulletin de la Societe Geologique deFrance, 178, 497–502.
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
The phylogeny of Tetanurae 279
Buffetaut, E., Bulow, M., Gheerbrant, E., Jaeger, J.-J., Martin,M., Mazin, J.-M., Milsent, C. & Rioult, M. 1985. Zonationbiostratigraphique et nouveaux restes de Vertebres dans les“Sables de Glos” (Oxfordien superieur, Normandie). ComptesRendus de l’Academie des Sciences a Paris, Serie 2, 300,929–932.
Buffetaut, E., Pennetier, G. & Pennetier, E. 1991a. Un fragmentde machoire de Megalosaurus dans le Callovien superier desVaches Noires (Calvados, France). Revue de Paleobiologie,10, 379–387.
Buffetaut, E., Cuny, G. & Le Loeuff, J. 1991. French dinosaurs:the best record in Europe? Modern Geology, 16, 17–42.
Buffetaut, E., Suteethorn, V. & Tong, H. 1996. The earliestknown tyrannosaur from the Lower Cretaceous of Thailand,Nature, 381, 689–691.
Buffetaut, E., Suteethorn, V. & Tong, H. 2004. Asian spinosaurconfirmed. In M. Evans & R. Forrest (eds) 52nd AnnualSymposium of Vertebrate Palaeontology and ComparativeAnatomy, Abstract Volume.
Buffetaut, E., Suteethorn, V., Le Loeuff, J., Khansubha, S.,Tong, H. & Wongko, K. 2005. The dinosaur fauna from theKhok Kruat Formation (Early Cretaceous) of Thailand. Pp.575–581 in Proceedings of the International Conference onGeology, Geotechnology and Mineral Resources of Indochina(GEOINDO 2005), 28–30 November 2005. Khon Kaen, Thai-land.
Bunzel, E. v. 1871. Die Reptilfauna der Gosau-Formation inder Neuen Welt bei Wiener-Neustadt. Abhandlungen derKaiserlich-Koniglichen Geologischen Reichsanstalt, 5, 1–18.
Buscalioni, A. D. & Sanz, J. L. 1984. Los Arcosaurios (Reptilia)del Jurasico superior – Cretacico inferior de Galve (Teruel,Espana). Teruel, 71, 9–30.
Bybee, P. J., Lee, A. H. & Lamm, E.-T. 2006. Sizing the Jurassictheropod dinosaur Allosaurus: assessing growth strategy andevolution of ontogenetic scaling of limbs. Journal of Morphol-ogy, 267, 347–359.
Calvo, J. O., Porfiri, J. D., Veralli, C. & Novas, F. E. 2002.Megaraptor namunhuaiquii (Novas, 1998), a new light aboutits phylogenetic relationships. Pp. 20–21 in 1◦ Congreso Lati-noamericano de Paleontologıa de Vertebrados, Resumenes.Santiago, Chile.
Calvo, J. O., Porfiri, J. D., Veralli, C., Novas, F. E. & Poblete,F. 2004. Phylogenetic status of Megaraptor namunhuaiquiiNovas based on a new specimen from Neuquen, Patagonia,Argentina. Ameghiniana, 41, 565–575.
Canale, J. I., Scanferla, C. A., Agnolin, F. L. & Novas, F. E.2009. New carnivorous dinosaur from the Late Cretaceousof NW Patagonia and the evolution of abelisaurid theropods.Naturwissenschaften, 96, 409–414.
Candeiro, C. R. A., Abranches, C. T., Abrantes, E. A., Avilla, L.S., Martins, V. C., Moreira, A. L., Torres, S. R. & Bergqvist,L. P. 2004. Dinosaurs remains from western Sao Paulo state,Brazil (Bauru Basin, Adamantina Formation, Upper Creta-ceous). Journal of South American Earth Sciences, 18, 1–10.
Candeiro, C. R. A., Martinelli, A. G., Avilla, L. S. &Rich, T. H. 2006. Tetrapods from the Upper Cretaceous(Turonian–Maastrichtian) Bauru Group of Brazil: a reap-praisal. Cretaceous Research, 27, 923–946.
Canudo, J. I. & Ruiz-Omenaca, J. I. 2003. Los restos directos dedinosaurios teropodos (excluyendo Aves) en Espana. Cienciasde la Tierra, 26, 347–373.
Canudo, J. I., Gasulla, J. M., Ortega, F. & Ruiz-Omenaca,J. I. 2004a. Presencia de Baryonychinae (Theropoda) en elAptiense inferior (Cretacico Inferior) de Laurasia: CanteraMas de la Parreta, Formacion Arcillas de Morella (Morella,Castellon). Pp. 11–12 in III Jornadas Internacionales sobre
Paleontologıa de Dinosaurios y su Entorno. Salas de losInfantes, Spain.
Canudo, J. I., Salgado, L., Barco, J. L., Bolatti, R. & Ruiz-Omenaca, J. I. 2004b. Dientes de dinosaurios teropodos ysauropodos de la Formacion Cerro Lisandro (Cenomaniensesuperior-Turoniense inferior, Cretacico superior) en RıoNegro (Argentina). Geo-Temas, 6, 31–34.
Canudo, J. I., M., Barco, J. L., Cuenca-Bescos, G. & Ruiz-Omenaca, J. I. 2005. Los dinosaurios de la FormacionVillar del Arzobispo (Titonico medio–Berriasiense inferior)en Galve (Teruel). Geogaceta, 38, 39–42.
Canudo, J. I., Filippi, L. S., Salgado, L., Garrido, A. C., Cerda,I., Garcıa, R. A. & Otero, A. 2007. Nuevos datos sobrelos titanosaurios (Dinosauria, Sauropoda) de la FormacionPlottier (Coniaciense–Santoniense) en Rincon de los Sauces(Neuquen, Patagonia, Argentina). Pp. 81–84 in E. Dıaz-Martınez & I. Rabano (eds) 4th European Meeting on thePalaeontology and Stratigraphy of Latin America, Madrid.Cuadernos del Museo Geominero, 8, Instituto Geologico yMinero de Espana, Madrid.
Canudo, J. I., Gasulla, J. M., Gomez-Fernandez, D., Ortega,F., Sanz, J. L. & Yague, P. 2008. Primer evidencia de dientesaislados atribuidos a Spinosauridae (Theropoda) en al Aptianoinferior (Cretacico Inferior) de Europa: Formacion Arcillas seMorella (Espana). Ameghiniana, 45, 649–662.
Carrano, M. T. 1999. What, if anything, is a cursor? Categoriesversus continua for determining locomotor habit in mammalsand dinosaurs. Journal of Zoology, 247, 29–42.
Carrano, M. T. 2006. Body-size evolution in the Dinosauria.Pp. 225–268 in M. T. Carrano, T. J. Gaudin, R. W. Blob &J. R. Wible (eds) Amniote Paleobiology: Perspectives on theEvolution of Mammals, Birds, and Reptiles. University ofChicago Press, Chicago.
Carrano, M. T. 2007. The appendicular skeleton of Majun-gasaurus crenatissimus (Theropoda: Abelisauridae) from theLate Cretaceous of Madagascar. Society of Vertebrate Paleon-tology Memoir, 8, 163–179.
Carrano, M. T. & Hutchinson J. R. 2002. The pelvic andhind limb musculature of Tyrannosaurus rex (Dinosauria:Theropoda). Journal of Morphology, 253, 207–228.
Carrano, M. T. & Sampson, S. D. 2004. A review of coelo-physoids (Dinosauria: Theropoda) from the Early Jurassicof Europe, with comments on the late history of the Coelo-physoidea. Neues Jahrbuch fur Geologie und PalaontologieMonatshefte, 2004, 537–558.
Carrano, M. T. & Sampson, S. D. 2008. The phylogeny ofCeratosauria (Dinosauria: Theropoda). Journal of SystematicPalaeontology, 6, 183–236.
Carrano, M. T., Sampson, S. D. & Forster, C. A. 2002. Theosteology of Masiakasaurus knopfleri, a small abelisauroid(Dinosauria: Theropoda) from the Late Cretaceous ofMadagascar. Journal of Vertebrate Paleontology, 22, 510–534.
Carrano, M. T., Hutchinson, J. R. & Sampson, S. D. 2005. Newinformation on Segisaurus halli, a small theropod dinosaurfrom the Early Jurassic of Arizona. Journal of VertebratePaleontology, 25, 835–849.
Carrano, M. T., Barrett, P. M. & Wilson, J. A. 2010. Thehistory of dinosaur collecting in central India, 1818–1947.Pp. 161–173 in R. Moody, E. Buffetaut, D. Naish & D. M.Martill (eds) Dinosaurs and Other Extinct Saurians: A Histor-ical Perspective. Geological Society Special Paper, 343. TheGeological Society, London.
Carrano, M. T., Loewen, M. A. & Sertich, J. J. W. 2011. Newmaterials of Masiakasaurus knopfleri Sampson, Carrano, andForster, 2001 and implications for the morphology of the
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
280 M. T. Carrano et al.
Noasauridae (Theropoda: Ceratosauria). Smithsonian Contri-butions to Paleobiology, 95, 1–53.
Carroll, R. R. 1988. Vertebrate Paleontology and Evolution. W.H. Freeman and Company, New York, 698 pp.
Casanovas-Cladellas, M. L., Santafe-Llopis, J. V. & Sanz-Garcıa, J. L. 1988. La primera resta fossil d’un teropode(Saurischia, Dinosauria) en el Cretaci Superior de la Conca deTremp (Lleda, Espana). Paleontologia i Evolucio, 22, 77–81.
Chabli, S. 1986. Donnees nouvelles sur un “Dinosaurien” Juras-sique moyen du Maroc: Megalosaurus mersensis Lapparent1955, et sur les Megalosaurides en general. Pp. 65–72 inLes Dinosauries de la Chine a la France, Colloque Inter-national de Paleontologie, Toulouse, France, 2–6 Septembre1985. Museum d’Histoire Naturelle de Toulouse, Toulouse.
Charig, A. J. & Milner, A. C. 1986. Baryonyx, a remarkable newtheropod dinosaur. Nature, 324, 359–361.
Charig, A. J. & Milner, A. C. 1990. The systematic position ofBaryonyx walkeri, in the light of Gauthier’s reclassification ofthe Theropoda. Pp. 127–140 in K. Carpenter & P. J. Currie(eds) Dinosaur Systematics: Approaches and Perspectives.Cambridge University Press, Cambridge.
Charig, A. J. & Milner, A. C. 1997. Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of theNatural History Museum, Geology Series, 53, 11–70.
Chure, D. J. 1995. A reassessment of the gigantic theropodSaurophagus maximus from the Morrison Formation (UpperJurassic) of Oklahoma, USA. Pp. 103–106 in A. Sun & Y.Wang (eds) Sixth Symposium on Mesozoic Terrestrial Ecosys-tems and Biota, Short Papers. China Ocean Press, Beijing.
Chure, D. J. 1998. A reassessment of the Australian Allosaurusand its implications for the Australian refugium concept. Jour-nal of Vertebrate Paleontology, 18(3, supplement), 34A.
Chure, D. J. 2000. Observations on the morphology and pathol-ogy of the gastral basket of Allosaurus, based on a new spec-imen from Dinosaur National Monument. Oryctos, 3, 29–37.
Chure, D. J. 2001a. A New Species of Allosaurus fromthe Morrison Formation of Dinosaur National Monument(Utah–Colorado) and a Revision of the Theropod FamilyAllosauridae. Unpublished PhD Thesis, Columbia University,964 pp.
Chure, D. J. 2001b. On the type and referred material of Laelapstrihedrodon Cope 1877 (Dinosauria: Theropoda). Pp. 10–18in D. H. Tanke & D. Carpenter (eds) Mesozoic Vertebrate Life.Indiana University Press, Bloomington.
Chure, D. J. & Madsen, J. H., Jr 1996. Variation in aspects ofthe tympanic pneumatic system in a population of Allosaurusfragilis from the Morrison Formation (Upper Jurassic). Jour-nal of Vertebrate Paleontology, 16, 63–66.
Chure, D. J., Manabe, M., Tanimoto, M. & Tomida, Y. 1999. Anunusual theropod tooth from the Mifune Group (Late Ceno-manian to Early Turonian), Kumamoto, Japan. Pp. 291–296in Y. Tomida, T. H. Rich & P. Vickers-Rich (eds) Proceedingsof the Second Gondwanan Dinosaur Symposium. NationalScience Museum Monographs, 15, National Science Museum,Tokyo.
Chure, D. J., Britt, B. B., Foster, J. R., Madsen, J. H., & Miles,C. A. 2000. New records of Ceratosaurus, Torvosaurus,Coelurus, and Stokesosaurus and their implication for thero-pod community structure and biozonation in the Late Jurassicof the Western Interior. Journal of Vertebrate Paleontology,20 (3, supplement), 35A.
Chure, D. J., Litwin, R. J., Hasiotis, S. T., Evanoff, E. &Carpenter, K. 2006. The fauna and flora of the MorrisonFormation: 2006. Pp. 233–249 in J. R. Foster & S. G. Lucas(eds) Paleontology and Geology of the Upper Jurassic Morri-
son Formation. New Mexico Museum of Natural History andScience Bulletin, 36, New Mexico Museum of Natural Historyand Science, Albuquerque.
Colbert, E. H. 1964. Relationships of the saurischian dinosaurs.American Museum Novitates, 2181, 1–24.
Colbert, E. H. & Russell, D. A. 1969. The small Creta-ceous dinosaur Dromaeosaurus. American Museum Novi-tates, 2380, 1–49.
Cope, E. D. 1867. An account of the extinct reptiles whichapproached the birds. Proceedings of the Academy of NaturalSciences of Philadelphia, 19, 234–235.
Cope, E. D. 1877. On a carnivorous dinosaur from the DakotaBeds of Colorado. Bulletin of the United States Geologicaland Geographical Survey, 3, 805–806.
Cope, E. D. 1878. The affinities of the Dinosauria. The AmericanNaturalist, 12, 57–58.
Cope, E. D. 1887a. The dinosaurian genus Coelurus. The Ameri-can Naturalist, 21, 367–369.
Cope, E. D. 1887b. A contribution to the history of the Vertebrataof the Trias of North America. Proceedings of the AmericanPhilosophical Society, 24, 209–228.
Cope, E. D. 1889. On a new genus of Triassic Dinosauria. TheAmerican Naturalist, 23, 626.
Cope, E. D. 1892. Skull of the dinosaurian Laelaps incrassatusCope. Proceedings of the American Philosophical Society,30, 240–245.
Corbella, H., Novas, F. E., Apesteguıa, S. & Leanza, H. A.2004. First fission-track age for the dinosaur-bearing NeuquenGroup (Upper Cretaceous), Neuquen Basin, Argentina.Revista del Museo Argentino de Ciencias Naturales, nuevoserie, 6, 227–232.
Coria, R. A. & Currie, P. J. 2002. The braincase of Gigan-otosaurus carolinii (Dinosauria, Theropoda) from the UpperCretaceous of Argentina. Journal of Vertebrate Paleontology,22, 802–811.
Coria, R. A. & Currie, P. J. 2006. A new carcharodon-tosaurid (Dinosauria, Theropoda) from the Upper Cretaceousof Argentina. Geodiversitas, 28, 71–118.
Coria, R. A. & Salgado, L. 1995. A new giant carnivorousdinosaur from the Cretaceous of Patagonia. Nature, 377,224–226.
Coria, R. A. & Salgado, L. 2000. A basal Abelisauria Novas 1992(Theropoda–Ceratosauria) from the Cretaceous of Patagonia,Argentina. GAIA, 15, 89–102.
Coria, R. A. & Salgado, L. 2005. Last Patagonian non-aviantheropods. Pp. 153–160 in K. Carpenter (ed.) The CarnivorousDinosaurs. Indiana University Press, Bloomington.
Coria, R. A., Chiappe, L. M. & Dingus, L. 2002. A new closerelative of Carnotaurus sastrei Bonaparte 1985 (Theropoda:Abelisauridae) from the Late Cretaceous of Patagonia. Jour-nal of Vertebrate Paleontology, 22, 460–465.
Currie, P. J. 1995. Phylogeny and systematics of theropods(Dinosauria). Journal of Vertebrate Paleontology, 15(3,supplement), 25A.
Currie, P. J. & Azuma, Y. 2006. New specimens, including agrowth series, of Fukuiraptor (Dinosauria, Theropoda) fromthe Lower Cretaceous Kitadani Quarry of Japan. Journal ofthe Paleontological Society of Korea, 22, 173–193.
Currie, P. J. & Carpenter, K. 2000. A new specimen ofAcrocanthosaurus atokensis (Theropoda, Dinosauria) fromthe Lower Cretaceous Antlers Formation (Lower Creta-ceous, Aptian) of Oklahoma, USA. Geodiversitas, 22, 207–246.
Currie, P. J. & Zhao, X.-J. 1994. A new carnosaur (Dinosauria,Theropoda) from the Jurassic of Xinjiang, People’s Republic
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
The phylogeny of Tetanurae 281
of China. Canadian Journal of Earth Sciences, 30 [for 1993],2037–2081.
Currie, P. J., Hurum, J. H. & Sabath, K. 2003. Skull structureand evolution in tyrannosaurid dinosaurs. Acta Palaeontolog-ica Polonica, 48, 227–234.
Cuvier, G. 1808. Sur les ossemens fossiles de crocodiles, et partic-ulierement sur ceux des environs du Havre et de Honfleur, avecdes remarques sur les squelettes des sauriens de la Thuringe.Annales du Museum d’Histoire Naturelle, 12, 73–110.
Dal Sasso, C. 2003. Dinosaurs of Italy. Comptes Rendus Palevol,2, 46–66.
Dal Sasso, C. & Brillante, G. 2001. Dinosauri Italiani. Marsilio,Venezia, 256 pp.
Dal Sasso, C., Maganuco, S., Buffetaut, E. & Mendez, M.A. 2005. New information on the skull of the enigmaticSpinosaurus, with remarks on its size and affinities. Journalof Vertebrate Paleontology, 25, 888–896.
Dames, W. 1884. Megalosaurus dunkeri. SitzungberichteGesellschaft Naturforschender Freunde zu Berlin, 1884,186–188.
Day, J. J. & Barrett, P. M. 2004. Material referred to Mega-losaurus (Dinosauria: Theropoda) from the Middle Juras-sic of Stonesfield, Oxfordshire, England: one taxon or two?Proceedings of the Geologists’ Association, 115, 359–366.
del Corro, G. 1966. Un nuevo Dinosaurio carnivoro del Chubut.Communicaciones del Museo Argentino de Ciencias Natu-rales “Bernardino Rivadavia” e Institutio Nacional de Inves-tigacion de las Ciencias Naturales: Paleontologia, 1, 1–4.
del Corro, G. 1974. Un nuevo Megalosaurio (Carnosaurio)del Cretacico de Chubut (Argentina). Communicaciones delMuseo Argentino de Ciencias Naturales “Bernardino Riva-davia” e Institutio Nacional de Investigacion de las CienciasNaturales: Paleontologia, 1, 37–44.
del Corro, G. 1975. Un nuevo sauropodo del Cretacico Supe-rior. Pp. 229–240 in Actas del Primer Congreso Argentino dePaleontologia y Bioestratigrafia. Asociacion PaleontologicaArgentina, Tucuman.
Delair, J. B. 1973. The dinosaurs of Wiltshire. Wiltshire Archae-ological and Natural History Magazine, 68, 1–7.
Delair, J. B. 1982. Multiple dinosaur trackways from the Isleof Purbeck. Proceedings of the Dorset Natural History andArchaeological Society, 102, 65–67.
Deperet, C. 1896. Note sur les dinosauriens sauropodes &theropodes du Cretace superieur de Madagascar. Bulletinde la Societe Geologique de France, 3e Serie, 24, 176–194.
Deperet, C. & Savornin, J. 1925. Sur la decouverte d’unefaune de Vertebres albiens a Timimoun (Sahara occidental).Comptes Rendus Hebdomadaires des Seances de l’Academiedes Sciences a Paris, 181, 1108–1111.
Deperet, C. & Savornin, J. 1928. La faune de Reptiles et dePoissons albiens de Timimoun (Sahara algerien). Bulletin dela Societe geologique de France, 4e Serie, 27 [for 1927],257–265.
Dingus, L., Clarke, J. A., Scott, G. R., Swisher, C. C., III,Chiappe, L. M. & Coria, R. A. 2000. Stratigraphy andmagnetostratigraphic/faunal constraints for the age of sauro-pod embryo-bearing rocks in the Neuquen Group (Late Creta-ceous, Neuquen Province, Argentina). American MuseumNovitates, 3290, 1–11.
Dollo, L. 1883. Note sur les restes de dinosauriens rencontres dansle Cretace superieur de la Belgique. Bulletin du Musee Royald’Histoire Naturelle de Belgique, 2, 205–221.
Dong, Z. 1973. [Dinosaurs from Wuerho]. Memoirs of the Instituteof Vertebrate Paleontology and Paleoanthropology Academ-ica Sinica, 11, 45–52. [In Chinese].
Dong, Z. 1984. [A new theropod dinosaur from the Middle Juras-sic of Sichuan Basin]. Vertebrata PalAsiatica, 22, 213–218[In Chinese].
Dong, Z. 1985. [A new Mid-Jurassic theropod (Gasosaurusconstructus gen. et sp. nov.) from Dashanpu, Zigong, SichuanProvince, China]. Vertebrata PalAsiatica, 23, 79–83 [InChinese].
Dong, Z. 1992. Dinosaurian Faunas of China. China Ocean Press,Beijing, 188 pp.
Dong, Z., Chang, Y. H., Li, X. M. & Zhou, S. W. 1978. [Note onthe new carnosaur (Yangchuanosaurus shangyouensis gen. etsp. nov.) from the Jurassic of Yangchuan District, SzechuanProvince.] Kexue Tongbao, 5, 302–304 [In Chinese].
Dong, Z., Zhou, S. W. & Zhang, H. 1983. [Dinosaurs from theJurassic of Sichuan]. Palaeontologica Sinica, New Series C,23 (Whole Number 162), 1–136 [In Chinese].
Downs, A. 2000. Coelophysis bauri and Syntarsus rhodesiensiscompared, with comments on the preparation and preservationof fossils from the Ghost Ranch Coelophysis Quarry. Pp.33–37 in S. G. Lucas & A. B. Heckert (eds) Dinosaurs ofNew Mexico. New Mexico Museum of Natural History andScience Bulletin, 17.
Eddy, D. R. & Clarke, J. A. 2011. New information on the cranialanatomy of Acrocanthosaurus atokensis and its implicationsfor the phylogeny of Allosauroidea (Dinosauria: Theropoda).PLoS ONE, 6, e17932 (doi:10.1371/journal.pone.0017932).
El-Zouki, A. Y. 1980. Stratigraphy and lithofacies of the continen-tal clastics (Upper Jurassic and Lower Cretaceous) of JabalNafusah, NW Libya. Pp. 393–417 in M. J. Salem & M. T.Busrewil (eds) The Geology of Libya. Second Symposium onthe Geology of Libya, Held at Tripoli, September 16–21, 1978.Academic Press, London.
Eudes-Deslongchamps, J. A. 1837. Memoire sur le Poekilopleu-ron bucklandi, grande saurien fossile, intermediare entre lescrocodiles et les lezards, decouvert dans les carrieres de laMaladrerie, pres Caen, au mois de juillet 1835. Memoires dela Societe Linneenne Normandie, 6, 1–114.
Ezcurra, M. D. 2006. A review of the systematic position of thedinosauriform archosaur Eucoelophysis baldwini Sullivan &Lucas, 1999 from the Upper Triassic of New Mexico, USA.Geodiversitas, 28, 649–684.
Ezcurra, M. D. & Novas, F. E. 2007. Phylogenetic relation-ships of the Triassic theropod Zupaysaurus rougieri from NWArgentina. Historical Biology, 19, 35–72.
Farlow, J. O., Smith, M. B. & Robinson, J. M. 1995. Bodymass, bone “strength indicator,” and cursorial potential ofTyrannosaurus rex. Journal of Vertebrate Paleontology, 15,713–725.
Fitzinger, L. 1843. Systema Reptilium. Fasciculus Primus.Amblyglossae. Apud Braumuller and Seidel Bibliopolas,Vienna, 106 pp.
Forster, C. A. 1999. Gondwanan dinosaur evolution and biogeo-graphic analysis. Journal of African Earth Sciences, 28,169–185.
Forster, C. A., Farke, A. A., McCartney, J. A., De Klerk, W.J. & Ross, C. F. 2009. A “basal” tetanuran from the LowerCretaceous Kirkwood Formation of South Africa. Journal ofVertebrate Paleontology, 29, 283–285.
Foster, J. R. 2003. Paleoecological analysis of the vertebrate faunaof the Morrison Formation (Upper Jurassic), Rocky Mountainregion, U.S.A. New Mexico Museum of Natural History andScience Bulletin, 23, 1–95.
Foster, J. R. & Chure, D. J. 2000. An ilium of a juvenile Stoke-sosaurus (Dinosauria, Theropoda) from the Morrison Forma-tion (Upper Jurassic: Kimmeridgian), Meade County, SouthDakota. Brigham Young University Geology Studies, 45, 5–10.
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
282 M. T. Carrano et al.
Fraser, N. C., Padian, K., Walkden, G. M. & Davis, A. L.M. 2002. Basal dinosauriform remains from Britain and thediagnosis of the Dinosauria. Palaeontology, 45, 79–95.
Galton, P. M. 1976. Iliosuchus, Jurassic dinosaur from Oxford-shire and Utah. Palaeontology, 19, 587–589.
Galton, P. M. 1998. Saurischian dinosaurs from theUpper Triassic of England: Camelotia (Prosauropoda,Melanorosauridae) and Avalonianus (Theropoda,?Carnosauria). Palaeontographica Abteilung A, 250, 155–172.
Galton, P. M. 2005. Bones of large dinosaurs (Prosauropoda andStegosauria) from the Rhaetic Bone Bed (Upper Triassic) ofAust Cliff, southwest England. Revue de Paleobiologie, 24,51–74.
Galton, P. M. & Jensen, J. A. 1979. A new large theropoddinosaur from the Upper Jurassic of Colorado. Brigham YoungUniversity Geology Studies, 26, 1–12.
Gao, Y. 1992. [Yangchuanosaurus hepingensis – a new species ofcarnosaur from Zigong, Sichuan]. Vertebrata PalAsiatica, 30,313–324 [In Chinese].
Gao, Y. 1993. [A new species of Szechuanosaurus from theMiddle Jurassic of Dashanpu, Zigong, Sichuan]. VertebrataPalAsiatica, 31, 308–314 [In Chinese].
Gao, Y. 1999. [Complete carnosaur skeleton from Zigong,Sichuan]. Zigong Dinosaur Museum, Zigong. 100 pp. [inChinese].
Gasca, J. M., Moreno-Azanza, M. & Canudo, J. I. 2008. Dientesde dinosaurios teropodos espinosauridos de la Formacion ElCastellar (Cretacico Inferior, Teruel). Palaeontologica Nova,8, 233–234.
Gauthier, J. A. 1986. Saurischian monophyly and the origin ofbirds. Pp. 1–47 in K. Padian (ed.) The Origin of Birds and theEvolution of Flight. Memoirs of the California Academy ofSciences, 8.
Gervais, P. 1853. Observations relatives aux Reptiles fossiles deFrance (deuxieme partie). Comptes Rendus de l’Academie desSciences a Paris, 36, 470–474.
Gervais, P. 1859. Zoologie et Paleontologie Francaise, 2me
Edition. Paris, Arthur Bertrand, 544 pp.Ghevariya, Z. G. 1988. Intertrappean dinosaur fossils from Anjar
area, Kachchh district, Gujarat. Current Science, 57, 248–251.Gilmore, C. W. 1920. Osteology of the carnivorous Dinosauria in
the United States National Museum, with special referenceto the genera Antrodemus (Allosaurus) and Ceratosaurus.Bulletin of the United States National Museum, 110, 1–154.
Goloboff, P. A., Farris, J. S. & Nixon, K. C. 2008. TNT, a freeprogram for phylogenetic analysis. Cladistics, 24, 774–786.
Gomez-Fernandez, D., Canudo, J. I. & Cano-Llop, V. 2007a.Descripcion de la cintura pelviana de un nuevo dinosaurioteropodo de la Formacion Morella (Aptiense inferior) en Valli-bona (Castellon, Espana). Pp. 71–72 in IV Jornadas Interna-cionales sobre Paleontologıa de Dinosaurios y su Entorno.Salas de los Infantes, Spain.
Gomez-Fernandez, D., Canudo, J. I. & Cano-Llop, V. 2007b.Cintura pelviana de un nuevo dinosaurio teropodo dela Formacion Morella (Aptiense inferior) en Vallibona(Castellon). Pp. 96–97 in J.C. Braga, A. Checa and M.Company (eds) XXIII Jornadas de la Sociedad Espanolade Paleontologıa. Instituto Geologico y Minero de Espanay Universidad de Granada, Caravaca de la Cruz.
Goodwin, M. B., Clemens, W. A., Hutchison, J. H., Wood, C.B., Zavada, M. S., Kemp, A., Duffin, C. J. & Schaff, C.R. 1999. Mesozoic continental vertebrates with associatedpalynostratigraphic dates from the northwestern Ethiopianplateau. Journal of Vertebrate Paleontology, 19, 728–741.
Greppin, J.-B. 1870. Description geologique du Jura Bernois etde quelques districts adjacents, compris dans la feuille VIIde l’Atlas Federal. Materiaux pour la carte geologique de LaSuisse, 8, 1–357.
Hammer, W. R. & Hickerson, W. J. 1994. A crested theropoddinosaur from Antarctica. Science, 264, 828–830.
Hammer, W. R. & Hickerson, W. J. 1999. Gondwana dinosaursfrom the Jurassic of Antarctica. Pp. 211–217 in Y. Tomida, T.H. Rich & P. A. Vickers-Rich (eds) Proceedings of the SecondGondwanan Dinosaur Symposium. National Science MuseumMonographs, Tokyo, 15.
Hanna, R. R. 2002. Multiple injury and infection in a sub-adulttheropod dinosaur Allosaurus fragilis with comparisons toallosaur pathology in the Cleveland-Lloyd Dinosaur QuarryCollection. Journal of Vertebrate Paleontology, 22, 76–90.
Harris, J. D. 1998a. A reanalysis of Acrocanthosaurus atokensis,its phylogenetic status, and paleobiogeographic implications,based on a new specimen from Texas. New Mexico Museumof Natural History and Science Bulletin, 13, 1–75.
Harris, J. D. 1998b. Large, Early Cretaceous theropods in NorthAmerica. Pp. 225–228 in S. G. Lucas, J. I. Kirkland & J. W.Estep (eds) Lower and Middle Cretaceous Terrestrial Ecosys-tems. New Mexico Museum of Natural History and ScienceBulletin, 14.
Hasegawa, Y., Murata, M., Wasada, K. & Manabe, M. 1992.The first carnosaur (Saurischia; Theropoda) from Japan: atooth from the Cenomanian Mifune Group of Kyushu. ScienceReports of the Yokohama National University, Section II, 39,41–49.
Hay, O. P. 1902. Bibliography and Catalogue of the Fossil Verte-brata of North America. Bulletin of the United States Geolog-ical Survey, 179, 1–868.
He, X. 1984. [The Vertebrate Fossils of Sichuan]. Sichuan Scien-tific and Technological Publishing House, Chengdu, 168 pp.[In Chinese].
Henry, M. J. 1876. L’Infralias dans la Franche-Comte. Memoiresde la Societe d’Emulation du Doubs, 4e Serie, 10, 287–476.
Hitchin, R. & Benton, M. J. 1997. Congruence between parsi-mony and stratigraphy: comparisons of three indices. Paleo-biology, 23, 20–33.
Hocknull, S. A., White, M. A, Tischler, T. R., Cook,A. G., Calleja, N. D., Sloan, T. & Elliott, D. A.2009. New Mid-Cretaceous (latest Albian) dinosaurs fromWinton, Queensland, Australia. PLoS One 4(7), e6190(doi:10.1371/journal.pone.0006190).
Holtz, T. R., Jr 1994a. The phylogenetic position of the Tyran-nosauridae: implications for theropod systematics. Journal ofPaleontology, 68, 1100–1117.
Holtz, T. R., Jr 1994b. The arctometatarsalian pes, anunusual structure of the metatarsus of Cretaceous Theropoda(Dinosauria: Saurischia). Journal of Vertebrate Paleontology,14, 480–519.
Holtz, T. R., Jr 2000. A new phylogeny of the carnivorousdinosaurs. GAIA, 15, 5–61.
Holtz, T. R., Jr 2004. Tyrannosauroidea. Pp. 111–136 in D.B. Weishampel, P. Dodson, & H. Osmolska (eds) TheDinosauria. Second edition. University of California Press,Berkeley.
Holtz, T. R., Jr, Molnar, R. E. & Currie, P. J. 2004. BasalTetanurae. Pp. 71–110 in D. B. Weishampel, P. Dodson & H.Osmolska (eds) The Dinosauria. Second edition. Universityof California Press, Berkeley.
Hone, D. W. E., Keesey, T. M., Pisani, D. & Purvis, A. 2005.Macroevolutionary trends in the Dinosauria: Cope’s Rule.Journal of Evolutionary Biology, 18, 587–595.
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
The phylogeny of Tetanurae 283
Hooijer, D. A. 1968. A Cretaceous dinosaur from the Syrian ArabRepublic. Koninklijke Nederlandse Akademie van Weten-schappen – Amsterdam, Proceedings Series B, 71, 150–152.
Howarth, M. K. 1980. Toarcian correlation chart. Pp. 53–73 in J.C. W. Cope, T. A. Getty, M. K. Howarth, N. Morton, & H. S.Torrens (eds) A Correlation of Jurassic Rocks in the BritishIsles. Part One: Introduction and Lower Jurassic. GeologicalSociety Special Report, 14.
Hu, S.-Y. 1963. [The carnivorous dinosaurian remains from Fusin,Liaoning]. Vertebrata PalAsiatica, 7, 174–176 [In Chinese].
Hu, S.-Y. 1964. [Carnosaurian remains from Alashan, InnerMongolia]. Vertebrata PalAsiatica, 8, 42–63 [In Chinese].
Hu, S. 1993. A new Theropoda (Dilophosaurus sinensis sp. nov.)from Yunnan, China. Vertebrata PalAsiatica, 31, 65–69.
Huelsenbeck, J. P. 1994. Comparing the stratigraphic record toestimates of phylogeny. Paleobiology, 20, 470–483.
Huene, F. von 1908. Die Dinosaurier der Europaischen Trias-formation, mit Berucksichtigung der aussereuropaischenVorkommnisse. Geologische und Palaeontologische Abhand-lungen, Supplement 1 (1), 1–419.
Huene, F. von. 1909. Skizze zu einer Systematik und Stammes-geschichte der Dinosaurier. Centralblatt fur Mineralogie,Geologie und Palaontologie, 1909, 12–22.
Huene, F. von 1914a. Das naturliche System der Saurischia.Centralblatt fur Mineralogie, Geologie und Palaontologie,1914, 154–158.
Huene, F. von 1914b. The dinosaurs not a natural order.The American Journal of Science, Series 4, 38, 145–146.
Huene, F. von 1914c. Beitrage zur Geschichte der Archosaurier.Geologie und Palaontologie Abhandlungen, 13(7), 1–56.
Huene, F. von 1914d. Saurischia and Ornithischia. GeologicalMagazine, 9, 444–445.
Huene, F. von 1920. Bemerkungen zur Systematik und Stam-megeschichte einiger Reptilian. Zeitschrift fur InduktiveAbstammungs- und Vererbungslehre, 24, 209–212.
Huene, F. von 1923. Carnivorous Saurischia in Europe since theTriassic. Bulletin of the Geological Society of America, 34,449–458.
Huene, F. von 1926a. The carnivorous Saurischia in the Juraand Cretaceous formations, principally in Europe. Revista delMuseo de La Plata, 29, 35–167.
Huene, F. von 1926b. On several known and unknown reptilesof the order Saurischia from England and France. Annals andMagazine of Natural History, Series 9, 17, 473–489.
Huene, F. von 1927. Sichtung der Grundlagen der jetzigenKenntnis der Sauropoden. Eclogae Geologica Helveticae, 20,444–470.
Huene, F. von. 1929. Kurze Ubersicht uber die Saurischia undihre naturlichen Zusammenhange. Palaontologische Zeit-schrift, 11, 269–273.
Huene, F. von 1932. Die fossile Reptil-Ordnung Saurischia, ihreEntwicklung und Geschichte. Monographien zur Geologieund Palaontologie, Series 1, 4, 1–361.
Huene, F. von 1956. Palaontologie und Phylogenie der NiederenTetrapoden. VEB Gustav Fischer Verlang, Jena, 716 pp.
Huene, F. von. 1959. Saurians in China and their relations.Vertebrata PalAsiatica, 3, 119–123.
Huene, F. von 1966. Ein Megalosauriden-Wirbel des Lias ausnorddeutschem Geschiebe. Neues Jahrbuch fur Geologie undPalaontologie, 1966, 318–319.
Hulke, J. W. 1879. Note on Poikilopleuron bucklandi of EudesDeslongchamps (pere), identifying it with Megalosaurusbucklandi. Quarterly Journal of the Geological Society ofLondon, 35, 233–238.
Hunt, A. P. & Lucas, S. G. 1991. Rioarribasaurus, a new name fora Late Triassic dinosaur from New Mexico (USA). Palaontol-ogische Zeitschrift, 65, 191–198.
Hutchinson, J. R. 2001. The evolution of femoral osteology andsoft tissues on the line to extant birds (Neornithes). ZoologicalJournal of the Linnean Society, 131, 169–197.
Hutt, S. & Newberry, P. 2004. An exceptional theropod verte-bra from the Wessex Formation (Lower Cretaceous), Isle ofWight, England. Proceedings of the Isle of Wight NaturalHistory and Archaeology Society, 20, 61–76.
Hutt, S., Martill, D. M. & Barker, M. J. 1996. The first Euro-pean allosaurid dinosaur (Lower Cretaceous, Wealden Group,England). Neues Jahrbuch fur Geologie und PalaontologieMonatshefte, 1996, 635–644.
Huxley, T. H. 1869. The classification of the Dinosauria, withobservations on the Dinosauria of the Trias. Quarterly Journalof the Geological Society, 26, 32–50.
Huxley, T. H. 1870. On the upper jaw of Megalosaurus. QuarterlyJournal of the Geological Society, 25 [for 1869], 311–314.
Infante, P., Canudo, J. I. & Ruiz-Omenaca, J. I. 2005. Primeraevidencia de dinosaurios teropodos en la Formacion Miram-bel (Barremiense inferior, Cretacico Inferior) en Castellote,Teruel. Geogaceta, 38, 31–34.
Irmis, R. B., Nesbitt, S. J., Padian, K., Smith, N. D., Turner,A. H., Woody, D. T. & Downs, A. 2007. A Late Triassicdinosauromorph assemblage from New Mexico and the riseof dinosaurs. Science, 317, 358–361.
Ivie, M. A., Slipinski, S. A. & Wegrzynowicz, P. 2001. Generichomonyms in the Coldiinae (Coleoptera: Zopheridae). InsectaMundi, 15, 63–64.
Jackson, J. F. 1931. A catalogue of Cretaceous fossils in theMuseum of Isle of Wight Geology, the Free Library, Sandown.Part II: Lower Cretaceous. Proceedings of the Isle of WightNatural History and Archaeological Society, 2 [for 1930],45–61.
Jaekel, O. v. 1910. Ueber einen neuen Belodonten ausdem Buntsandstein von Bernburg. Sitzungsberichte derGesellschaft Naturforschender Freunde zu Berlin, 5, 197–229.
Janensch, W. 1920. Uber Elaphrosaurus bambergi und die Mega-losaurier aus den Tendaguru-Schichten Deutsch-Ostafrikas.Sitzungsberichte der Gesellschaft Naturforschender Freundezu Berlin, 1920, 225–235.
Janensch, W. 1925. Die Coelurosaurier und Theropoden derTendaguru-Schichten Deutsch-Ostafrikas. Palaeontograph-ica Supplement, 7(1), 1–100.
Jensen, J. A. 1985. Uncompaghre dinosaur fauna: a prelim-inary report. The Great Basin Naturalist, 45, 710–720.
Kellner, A. W. A. & Campos, D. A. 1996. First Early Creta-ceous theropod dinosaur from Brazil with comments onSpinosauridae. Neues Jahrbuch fur Geologie und Palaontolo-gie, Abhandlungen, 199, 151–166.
Kiprianow, W. 1883. Studien uber die Fossilen Reptilien Russ-lands. IV. Theil. Ordnung Crocodilina Oppel. IndeterminirteFossile Reptilien. Memoires de l’Academie Imperiale desSciences de St.-Petersbourg, VIIe serie, 31, 1–29.
Knoll, F., Buffetaut, E. & Bulow, M. 1999. A theropod brain-case from the Jurassic of the Vaches Noires cliffs (Normandy,France): osteology and paleoneurology. Bulletin de la SocieteGeologique de France, 1999, 103–109.
Kobayashi, Y. & Lu, J.-C. 2003. A new ornithomimid dinosaurwith gregarious habits from the Late Cretaceous of China.Acta Palaeontologica Polonica, 48, 235–259.
Krause, D. W., Sampson, S. D., Carrano, M. T. & O’Connor,P. M. 2007. Overview of the history of discovery, taxonomy,
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
284 M. T. Carrano et al.
phylogeny, and biogeography of Majungasaurus crenatis-simus (Theropoda: Abelisauridae) from the Late Cretaceousof Madagascar. Society of Vertebrate Paleontology Memoir,8, 1–20.
Kuhn, O. 1939. Saurischia Fossilium Catalogus 1: Animalia. Pars87: Saurischia. W. Junk, The Hague, 124 pp.
Kuhn, O. 1965. Fossilium Catalogus 1: Animalia. Pars 109:Saurischia. Ysel Press, Deventer, 94 pp.
Kurzanov, S. M. 1989. O proiskhozhdenii i evolyutsiiinfraotryada dinozavrov Carnosauria. PaleontologicheskiiZhurnal, 1989(4), 3–14.
Kurzanov, S. M., Efimov, M. B. & Gubin, Y. M. 2003. Newarchosaurs from the Jurassic of Siberia and Mongolia. Pale-ontological Journal, 37(1), 53–57.
Kutoba K. 2005. Charophyte gyrogonites from the Lower Creta-ceous Kitadani Formation of the Tetori Group in the Takinami-gawa area, Katsuyama City, Fukui Prefecture, central Japan.Paleontological Research, 9, 203–213.
Lamanna, M. C., Smith, J. B., You, H., Holtz, T. R., Jr &Dodson, P. 1998. A reassessment of the Chinese theropoddinosaur Dilophosaurus sinensis. Journal of Vertebrate Pale-ontology, 18(3, supplement), 57A.
Langer, M. C. 2004. Basal Saurischia. Pp. 25–46 in D. B.Weishampel, P. Dodson & H. Osmolska (eds) The Dinosauria.Second edition. University of California Press, Berkeley.
Lapparent, A. F. de 1947. Les dinosauriens du Cretace Superieurdu midi de la France. Memoires de la Societe Geologique deFrance, 56, 1–54.
Lapparent, A. F. de 1955. Etude paleontologique des vertebresdu Jurassique d’El Mers (Moyen Atlas). Notes et Memoiresdu Service Geologique du Maroc, 124, 1–36.
Lapparent, A. F. de 1960. Les dinosauriens du “Continental Inter-calaire” du Sahara central. Memoires de la Societe Geologiquede France, nouvelle serie, 88A, 1–57.
Lapparent, A. F. de & Zbyszewski, G. 1957. Les dinosauriensdu Portugal. Services Geologiques du Portugal Memoires(nouvelle series), 2, 1–63.
Lavocat, R. 1955. Sur une portion de mandibule de theropodeprovenant du Cretace superieur de Madagascar. Bulletin duMuseum National d’Histoire Naturelle, Paris, 2e serie, 27(3),256–259.
Leanza, H. A., Apesteguıa, S., Novas, F. E. & de la Fuente,M. S. 2004. Cretaceous terrestrial beds from the NeuquenBasin (Argentina) and their tetrapod assemblages. CretaceousResearch, 25, 61–87.
Leidy, J. 1870. (Remarks on Poikilopleuron valens, Clidastesintermedius, Macrosaurus proriger, Baptemus wyomingensis,and Emys stevensonianus). Proceedings of the Academy ofNatural Sciences of Philadelphia, 22, 3–5.
Leidy, J. 1873. Contributions to the extinct vertebrate faunaof the western territories. Reports of the U.S. Geologi-cal Survey of the Territories. F.W. Hayden, Department ofthe Interior, Government Printing Office, Washington DC,358 pp.
Le Loeuff, J., Metais, E., Dutheil, D. B., Rubino, J. L., Buffe-taut, E., Lafont, F., Cavin, L., Moreau, F., Tong, H., Blan-pied, C. & Sbeta, A. 2010. An Early Cretaceous vertebrateassemblage from the Cabao Formation of NW Libya. Geolog-ical Magazine, 147, 750–759.
Lennier, G. 1870. Etudes Geologiques et Paleontologiques surl’Embouchure de la Seine et les Falaises de la Haute-Normandie. Imprimerie Eugene Costey, Havre, 245 pp.
Lennier, G. 1887. Etudes paleontologiques. Description desfossils du Cap de la Heve. Bulletin de la Societe Geologiquede Normandie, 12 [for 1886], 17–98.
Li, F., Peng, G., Yong, Y., Jiang, S. & Huang, D. 2009. A newcarnosaur from the Late Jurassic of Qianwei, Sichuan, China.Acta Geologica Sinica, 83, 1203–1210.
Lim, J.-D., Martin, L. D. & Baek, K. S. 2002. The firstmegalosaurid tooth from South Korea. Current Science, 82,326–328.
Lipka, T. R. 1998. The affinities of the enigmatic theropods of theArundel Clay facies (Aptian), Potomac Formation, AtlanticCoastal Plain of Maryland. Pp. 229–234 in S. G. Lucas, J.I. Kirkland & J. W. Estep (eds) Lower and Middle Creta-ceous Terrestrial Ecosystems. New Mexico Museum of Natu-ral History and Science Bulletin, 14.
Loewen, M. A. 2009. Variation in the Late Jurassic TheropodDinosaur Allosaurus: Ontogenetic, Functional, and Taxo-nomic Implications. Unpublished PhD Thesis, University ofUtah, 326 pp.
Long, J. A. 1995. A theropod dinosaur bone from the Late Creta-ceous Molecap Formation, Western Australia. Records of theWestern Australian Museum, 17, 143–146.
Long, J. A. & Cruickshank, A. R. I. 1996. First record of an EarlyCretaceous theropod dinosaur bone from Western Australia.Records of the Western Australian Museum, 18, 219–222.
Lu, J. & Hu, P. 1998. [Dinosaur remains from Datong Suburb,Shanxi Province]. Vertebrata PalAsiatica, 36, 252–256 [InChinese].
Lydekker, R. 1888. A Catalogue of the Fossil Reptilia andAmphibia in the British Museum. Part I. Containing theOrders Ornithosauria, Crocodilia, Dinosauria, Squamata,Rhynchocephalia, and Proterosauria. British Museum (Natu-ral History), London, 309 pp.
Lydekker, R. 1889. On the remains and affinities of five generaof Mesozoic reptiles. Quarterly Journal of the GeologicalSociety of London, 45, 41–59.
Lydekker, R. 1890a. Catalogue of the Fossil Reptilia andAmphibia in the British Museum. Part IV. Containing theOrder Anomodonia, Ecaudata, Caudata, Labyrinthodonta;and Supplement. British Museum (Natural History), London,295 pp.
Lydekker, R. 1890b. Vertebrate remains from Nagpur District.Records of the Geological Survey of India, 23, 20–24.
Maddison, W. P. & Maddison, D. R. 1992. MacClade: Analysisof Phylogeny and Character Evolution. Sinauer, Sunderland,Massachusetts.
Maddison, W. P. & Maddison, D. R. 2009. Mesquite: A ModularSystem for Evolutionary Analysis. Version 2.72 Build 527.
Madsen, J. H., Jr 1974. A new theropod dinosaur from the UpperJurassic of Utah. Journal of Paleontology, 48, 27–31.
Madsen, J. H., Jr 1976a. Allosaurus fragilis: a revised osteol-ogy. Utah Geological and Mineral Survey Bulletin, 109, 1–163.
Madsen, J. H., Jr 1976b. A second new theropod dinosaur fromthe Late Jurassic of east central Utah. Utah Geology, 3, 51–60.
Madsen, J. H., Jr & Welles, S. P. 2000. Ceratosaurus(Dinosauria, Theropoda): a revised osteology. Utah Geologi-cal Survey, Miscellaneous Publications, 00–2, 1–80.
Malafaia, E., Dantas, P., Ortega, F. & Escaso, F. 2007. Nuevosrestos de Allosaurus fragilis (Theropoda: Carnosauria)del yacimiento de Andres (Jurasico Superior; centro-oestede Portugal). Pp. 255–271 in Cantera Paleontologica.Diputacion Provincial de Cuenca, Cuenca.
Mantell, G. A. 1827. Illustrations of the Geology of Sussex: AGeneral View of the Geological Relations of the South-EasternPart of England, with Figures and Descriptions of the Fossilsof Tilgate Forest. Lupton Relfe, London, 92 pp.
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
The phylogeny of Tetanurae 285
Marsh, O. C. 1877. Notice of new dinosaurian reptiles from theJurassic formation. The American Journal of Science and Arts,Series 3, 14, 514–516.
Marsh, O. C. 1878. Notice of new dinosaurian reptiles. TheAmerican Journal of Science and Arts, Series 3, 15, 241–244.
Marsh, O. C. 1879. Principal characters of American Jurassicdinosaurs. Part II. The American Journal of Science and Arts,Series 3, 17, 86–92.
Marsh, O. C. 1882. Classification of the Dinosauria. The Ameri-can Journal of Science and Arts, Series 3, 23, 81–86.
Marsh, O. C. 1884. Principal characters of American Jurassicdinosaurs. Part VIII. The order Theropoda. The AmericanJournal of Science and Arts, Series 3, 27, 329–340.
Marsh, O. C. 1895. On the affinities and classification of thedinosaurian reptiles. The American Journal of Science andArts, Series 3, 50, 483–498.
Marsh, O. C. 1896. The dinosaurs of North America. UnitedStates Geological Survey, 16th Annual Report, 1894-95, 55,133–244.
Marshall, C. R. 1994. Confidence intervals on stratigraphicranges: partial relaxation of the assumption of randomlydistributed fossil horizons. Paleobiology, 20, 459–469.
Martill, D. M. & Hutt, S. 1996. Possible baryonychid dinosaurteeth from the Wessex Formation (Lower Cretaceous,Barremian) of the Isle of Wight, England. Proceedings ofthe Geologists’ Association, 107, 81–84.
Martill, D. M., Cruickshank, A. R. I., Frey, E. D., Small, P.G. & Clarke, M. 1996. A new crested maniraptoran dinosaurfrom the Santana Formation (Lower Cretaceous) of Brazil.Journal of the Geological Society, London, 153, 5–8.
Martinelli, A. G. & Forasiepi, A. M. 2004. Late Cretaceousvertebrates from Bajo de Santa Rosa (Allen Formation),Rıo Negro province, Argentina, with the description of anew sauropod dinosaur (Titanosauridae). Revista del MuseoArgentino de Ciencias Naturales, Nuevo Serie, 6, 257–305.
Mateus, O. 1998. Lourinhanosaurus antunesi, a new UpperJurassic allosauroid (Dinosauria: Theropoda) from Lourinha,Portugal. Memorias da Academia de Ciencias de Lisboa, 37,111–124.
Mateus, O. & Antunes, M. T. 2000. Torvosaurus sp. (Dinosauria:Theropoda) in the Late Jurassic of Portugal. Livro de Resumosde I Congreso Iberico de Paleontologia/XVI Jornadas de laSociedad Espanola de Paleontologıa, 115–117.
Mateus, O., Walen, A. & Antunes, M. T. 2006. The large thero-pod fauna of the Lourinha Formation (Portugal) and its simi-larity to that of the Morrison Formation, with a descriptionof a new species of Allosaurus. Pp. 123–129 in J. R. Foster& S. G. Lucas (eds) Paleontology and Geology of the UpperJurassic Morrison Formation. New Mexico Museum of Natu-ral History and Science Bulletin, 36.
Mateus, O., Araujo, R., Natario, C. & Castanhinha, R. 2011.A new specimen of the theropod dinosaur Baryonyx fromthe early Cretaceous of Portugal and taxonomic validity ofSuchosaurus. Zootaxa, 2827, 54–68.
Matthew, W. D. 1915. Dinosaurs, with Special Reference to theAmerican Museum Collections. American Museum of NaturalHistory, New York, 162 pp.
Medeiros, M. A. 2005. Spinosaurid teeth variation in mid-Cretaceous Alcantara Formation, Maranhao state, Brazil.Resumos do XIX Congresso Brasileiro de Paleontologia eVI Congresso Latino-Americano de Paleontologia.
Medeiros, M. A. 2006. Large theropod teeth from the Eocenoma-nian of northeastern Brazil and the occurrence of Spinosauri-dae. Revista Brasileira de Paleontologia, 9, 333–338.
Medeiros, M. A. & Schultz, C. L. 2002. A fauna dinossaurianada “Laje do Coringa”, Cretaceo Medio do nordeste do Brasil.Arquivos do Museu Nacional, Rio de Janeiro, 60, 155–162.
Meyer, H. von 1832. Palaologica zur Geschichte der Erde. Sieg-mund Schmerber Verlag, Frankfurt-am-Main, 560 pp.
Meyer, C. A. & Thuring, B. 2003. Dinosaurs of Switzerland.Comptes Rendus Palevol, 2, 103–117.
Milner, A. C., Buffetaut, E. & Suteethorn, V. 2007. A tall-spinedspinosaurid theropod from Thailand and the biogeography ofspinosaurs. Journal of Vertebrate Paleontology, 27(3, supple-ment), 118A.
Molnar, R. E. 1990. Problematic Theropoda: “carnosaurs”. Pp.306–317 in D. B. Weishampel, P. Dodson & H. Osmolska (eds)The Dinosauria. University of California Press, Berkeley.
Molnar, R. E., Flannery, T. F. & Rich, T. H. 1981. An allosauridtheropod dinosaur from the Early Cretaceous of Victoria,Australia. Alcheringa, 5, 141–146.
Molnar, R. E., Flannery, T. F. & Rich, T. H. 1985. AussieAllosaurus after all. Journal of Paleontology, 59, 1511–1513.
Molnar, R. E., Kurzanov, S. M. & Dong, Z. 1990. Carnosauria.Pp. 169–209 in D. B. Weishampel, P. Dodson & H. Osmolska(eds) The Dinosauria. University of California Press, Berke-ley.
Molnar, R. E., Angriman, A. L. & Gasparini, Z. 1996. AnAntarctic Cretaceous theropod. Memoirs of the QueenslandMuseum, 39, 669–674.
Munster, G. Graf zu. 1836. Ueber die Korallenkalk das LinderBerges bei Hanover vorkommenden Ueberreste von Fischen,mit Beschreibung und Abbildung einiger neuen Arten. Pp.36–50 in G. Graf zu Munster & H. Wissman (eds) Beitragezur Petrefacten-Kunde, 7.
Naish, D. M. 2003. A definitive allosauroid (Dinosauria;Theropoda) from the Lower Cretaceous of East Sussex.Proceedings of the Geologists’ Association, 114, 319–326.
Naish, D. M. & Martill, D. M. 2007. Dinosaurs of Great Britainand the role of the Geological Society of London in theirdiscovery: basal Dinosauria and Saurischia. Journal of theGeological Society of London, 164, 493–510.
Naish, D. M., Hutt, S. & Martill, D. M. 2001. Saurischiandinosaurs 2: theropods. Palaeontological Association FieldGuide to Fossils, 10, 242–309.
Nesbitt, S. J., Smith, N. D., Irmis, R. B., Turner, A. H., Downs,A. & Norell, M. A. 2009. A complete skeleton of a Late Trias-sic saurischian and the early evolution of dinosaurs. Science,326, 1530–1533.
Nessov, L. A. 1991. A Late Jurassic labyrinthodont (Amphibia,Labyrinthodontia) along other relict groups of vertebratesfrom northern Fergana. Paleontological Journal, 24(3),83–90.
Nessov, L. A. 1995. Dinozavri severnoi Yevrazii: Novye dannye osostave kompleksov, ekologii I paleobiogeografii. Institute forScientific Research on the Earth’s Crust, St. Petersburg StateUniversity, St. Petersburg, 156 pp.
Newman, B. H. 1968. The Jurassic dinosaur Scelidosaurusharrisoni, Owen. Palaeontology, 11, 40–43.
Newton, E. T. 1899. On a megalosaurid jaw from the RhaeticBeds near Bridgend (Glamorganshire). Quarterly Journal ofthe Geological Society of London, 55, 89–96.
Nixon, K. C. 1999. The Parsimony Ratchet, a new method forrapid parsimony analysis. Cladistics, 15, 407–414.
Nopcsa, B. F. 1901. Synopsis und Abstammung der dinosaurier.Foldtani Kozlony, 31, 247–279.
Nopcsa, B. F. 1902. Notizen uber Cretacische Dinosaurier.Sitzungsberichte der Kaiserlichen Akademie der
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
286 M. T. Carrano et al.
Wissenschaften. Mathematisch-NaturwissenschaftlicheClasse, 111, 93–114.
Nopcsa, B. F. 1905. Notes on British dinosaurs. Part III: Strep-tospondylus. Geological Magazine, Decade 5, 2, 288–293.
Nopcsa, B. F. 1906. Zur Kenntnis des Genus Streptospondy-lus. Beitrage zur Palaontologie Osterreich-Ungarns und desOrients, 19, 59–83.
Nopcsa, B. F. 1923. Die Familien der Reptilien. Forschritte derGeologie und Palaeontologie, 2, 1–210.
Nopcsa, B. F. 1928. The genera of reptiles. Palaeobiologica, 1,163–188.
Novas, F. E. 1991a. Los Tyrannosauridae, gigantescos dinosaurioscelurosaurios del Cretacico tardio de Laurasia. Ameghiniana,28, 401.
Novas, F. E. 1991b. Relaciones filogeneticas de los dinosauriosteropodos ceratosaurios. Ameghiniana, 28, 401.
Novas, F. E. 1992. La evolucion de los dinosaurios carnivoros.Pp. 126–163 in J. L. Sanz & A. D. Buscalioni (eds) LosDinosaurios y Su Entorno Biotico: Actas del Segundo Cursode Paleontologıa en Cuenca. Instituto ‘Juan Valdez,’ Cuenca,Spain.
Novas, F. E. 1998. Megaraptor namunhuaiquii, gen. et sp. nov., alarge-clawed, Late Cretaceous theropod from Patagonia. Jour-nal of Vertebrate Paleontology, 18, 4–9.
Novas, F. E. 2008. Orkoraptor burkei nov. gen. et sp., a largetheropod from the Maastrichtian Pari Aike Formation, South-ern Patagonia, Argentina. Cretaceous Research, 29, 468–480.
Novas, F. E. & Agnolin., F. L. 2004. Unquillosaurus ceibaliPowell, a giant maniraptoran (Dinosauria, Theropoda) fromthe Late Cretaceous of Argentina. Revista del MuseoArgentino de Ciencias Naturales, nuevo serie, 6, 61–66.
Novas, F. E., de Valais, S., Vickers-Rich, P. A. & Rich,T. H. 2005. A large Cretaceous theropod from Patagonia,Argentina, and the evolution of carcharodontosaurids. Natur-wissenschaften, 92, 226–230.
Okazaki, Y. 1992. A new genus and species of carnivorousdinosaur from the Lower Cretaceous Kwanmon Group, north-ern Kyushu. Bulletin of the Kitakyushu Museum of NaturalHistory, 11, 87–90.
Olshevsky, G. 1991. A revision of the parainfraclass ArchosauriaCope, 1869, excluding the advanced Crocodylia. MesozoicMeanderings, 2, 1–196.
Olshevsky, G. 1995. African dinosaur discoveries. Science, 267,1750–1752.
Ortega, F., Escaso, F. & Sanz, J. L. 2010. A bizarre, humpedCarcharodontosauria (Theropoda) from the Lower Cretaceousof Spain. Nature, 467, 203–206.
Osborn, H. F. 1903. Ornitholestes hermanni, a new compsog-nathoid dinosaur from the Upper Jurassic. Bulletin of theAmerican Museum of Natural History, 19, 459–464.
Osborn, H. F. 1917. Skeletal adaptations of Ornitholestes,Struthiomimus, Tyrannosaurus. Bulletin of the AmericanMuseum of Natural History, 35, 733–771.
Osborn, H. F. 1931. Cope: Master Naturalist. Princeton Univer-sity Press, Princeton, 740 pp.
Ostrom, J. H. 1970. Stratigraphy and paleontology of the CloverlyFormation (Lower Cretaceous) of the Bighorn Basin area,Wyoming and Montana. Peabody Museum of Natural HistoryBulletin, 35, 1–234.
Ostrom, J. H. 1978. The osteology of Compsognathus longipesWagner. Zitteliana 4, 73–118.
Owen, R. 1840–1845. Odontography. Hippolyte Bailliere,London.
Owen, R. 1841. Description of a portion of the skeleton of theCetiosaurus, a gigantic extinct saurian reptile occurring in the
Oolitic formations of different portions of England. Proceed-ings of the Geological Society of London, 3(2), 457–462.
Owen, R. 1842. Report on British fossil reptiles, part II. Reportsof the British Association for the Advancement of Science, 11[for 1841], 60–204.
Owen, R. 1857. Monograph on the fossil Reptilia of the Wealdenand Purbeck Formations. Part III. Dinosauria (Megalosaurus).(Wealden). Monographs of the Palaeontological Society,London, 9(34, for 1855), 1–26.
Owen, R. 1858. Monograph on the fossil Reptilia of the Wealdenand Purbeck Formations. Part IV. Dinosauria (Hylaeosaurus).(Wealden). Monographs of the Palaeontological Society,London, 1856(43, for 1856), 8–26.
Owen, R. 1876. Monograph on the fossil Reptilia of theWealden and Purbeck formations. Supplement no. VII.Crocodilia (Poikilopleuron) and Dinosauria? (Chondros-teosaurus). (Wealden.). The Palaeontographical Society,London, 1876(136), 1–7.
Owen, R. 1883. On the skull of Megalosaurus. Quarterly Journalof the Geological Society of London, 39, 334–337.
Park, E.-J., Yank, S.-Y. & Currie, P. J. 2000. Early Cretaceousdinosaur teeth of Korea. Pp. 85–98 in Y.-N. Lee (ed.) 2000International Dinosaur Symposium for Kosong County inKorea. Paleontological Society of Korea Special Publication,4.
Paul, G. S. 1984. The archosaurs: a phylogenetic study. Pp.175–180 in W.-E. Reif & F. Westphal (eds) Third Symposiumon Mesozoic Terrestrial Ecosystems, Short Papers. AttemptoVerlag, Tubingen.
Paul, G. S. 1988a. Predatory Dinosaurs of the World. Simon &Schuster, New York, 464 pp.
Paul, G. S. 1988b. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and GreatOolite – Ornitholestes and Proceratosaurus – and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River– Deinonychus, Velociraptor and Saurornitholestes. Hunteria2(4), 1–9.
Paul, G. S. 2002. Dinosaurs of the Air. The Johns Hopkins Univer-sity Press, Baltimore, 472 pp.
Pereda Suberbiola, X., Torcida, F., Izquierdo, L. A.,Huerta, P., Montero, D. & Perez, G. 2003. Firstrebbachisaurid dinosaur (Sauropoda, Diplodocoidea) from theearly Cretaceous of Spain: palaeobiogeographical implica-tions. Bulletin de la Societe geologique de France, 2003, 471–479.
Perez-Moreno, B. P., Sanz, J. L. Sudre, J. & Sige, B. 1994.A theropod dinosaur from the Lower Cretaceous of southernFrance. Revue de Paleobiologie, Volume speciale, 7, 173–188.
Perez-Moreno, B. P., Chure, D. J., Pires, C., da Silva, C. M.,dos Santos, V. F., Dantas, P., Povoas, L., Cachao, M.,Sanz, J. L. & Carvalho, A. M. G. de 1999. On the pres-ence of Allosaurus fragilis (Theropoda: Carnosauria) in theUpper Jurassic of Portugal: first evidence of an interconti-nental dinosaur species. Journal of the Geological Society,London, 156, 449–452.
Peyer, K. 2006. A reconsideration of Compsognathus longipesfrom the Upper Tithonian of Canjuers, southeastern France.Journal of Vertebrate Paleontology, 26, 879–896.
Phillips, J. 1871. Geology of Oxford and the Valley of the Thames.Clarendon Press, Oxford, 523 pp.
Piveteau, J. 1923. L’arriere-crane d’un dinosaurien carnivore del’Oxfordien de Dives. Annales de Paleontologie, 12, 115–123.
Piveteau, J. 1955. Traite de Paleontologie. Tome V. L’Origine desVertebres, Leur Expansion Dans les Eaux Douces et le Milieu
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
The phylogeny of Tetanurae 287
Marin: Amphibiens, Reptiles, Oiseaux. Masson and Cie, Paris,1113 pp.
Porfiri, J. D. & Calvo, J. O. 2003. “Megaraptores” en lagoBarreales, Neuquen, Patagonia. XIX Jornadas Argentinas dePaleontologıa de Vertebrados. Ameghiniana, 40, 90R.
Powell, J. E. 1979. Sobre una asociacion de dinosaurios y otrasevidencias de vertebrados del Cretacico Superior de la regionde La Candelaria, Prov. de Salta, Argentina. Ameghiniana, 16,191–204.
Powell, H. P. 1987. A megalosaurid dinosaur jawbone from theKimmeridge Clay of the seabed of West Bay, Dorset. Proceed-ings of the Dorset Natural History and Archaeological Soci-ety, 109, 105–108.
Quenstedt, F. A. 1843. Das Flozgebirge Wurttembergs: mitbesonderer Rucksicht auf den Jura. Verlag der H. Laupp’schenBuchhandlung, Tubingen, 558 pp.
Quenstedt, F. A. 1852. Handbuch der Petrefaktenkunde. H.Laupp’schen Buchhandlung, Tubingen, 792 pp.
Quenstedt, F. A. 1858. Der Jura. Verlag der H. Laupp’schenBuchhandlung, Tubingen, 842 pp.
Raath, M. A. 1969. A new coelurosaurian dinosaur from theForest Sandstone of Rhodesia. Arnoldia (Rhodesia), 4(28),1–25.
Raath, M. A. 1977. The Anatomy of the Triassic Theropod Syntar-sus rhodesiensis (Saurischia: Podokesauridae) and a Consid-eration of Its Biology. Unpublished PhD Thesis, RhodesUniversity, 233 pp.
Raath, M. A. & McIntosh, J. S. 1987. Sauropod dinosaursfrom the central Zambezi Valley, Zimbabwe, and the age ofthe Kadzi Formation. South African Journal of Geology, 90,107–119.
Racey, A. & Goodall, J. G. S. 2009. Palynology and stratigraphyof the Mesozoic Khorat Group of NE Thailand. Pp. 68–81 inE. Buffetaut, G. Cuny, J. Le Loeuff & V. Suteethorn (eds) LatePalaeozoic and Mesozoic Ecosystems in SE Asia. GeologicalSociety of London Special Publication, 315.
Rauhut, O. W. M. 1995. Zur systematischen Stellung derafrikanischen Theropoden Carcharodontosaurus Stromer1931 und Bahariasaurus Stromer 1934. Berliner Geowis-senschaftliche Abhandlungen, E, 16, 357–375.
Rauhut, O. W. M. 1999. A dinosaur fauna from the LateCretaceous (Cenomanian) of northern Sudan. PalaeontologiaAfricana, 35, 61–84.
Rauhut O. W. M. 2000. The Interrelationships and Evolutionof Basal Theropods (Dinosauria, Saurischia). UnpublishedPh.D. Thesis, University of Bristol, 583 pp.
Rauhut, O. W. M. 2003. The interrelationships and evolution ofbasal theropod dinosaurs. Special Papers in Palaeontology,69, 1–213.
Rauhut, O. W. M. 2004. Braincase structure of the Middle Juras-sic theropod dinosaur Piatnitzkysaurus. Canadian Journal ofEarth Sciences, 41, 1109–1122.
Rauhut, O. W. M. 2005a. Osteology and relationships of a newtheropod dinosaur from the Middle Jurassic of Patagonia.Palaeontology, 48, 87–110.
Rauhut, O. W. M. 2005b. Post-cranial remains of ‘coelurosaurs’(Dinosauria, Theropoda) from the Late Jurassic of Tanzania.Geological Magazine, 142, 97–107.
Rauhut, O. W. M. 2007. A fragmentary theropod skull from theMiddle Jurassic of Patagonia. Ameghiniana, 44, 479–483.
Rauhut, O. W. M. & Fechner, R. 2005. Early development of thefacial region in a non-avian theropod dinosaur. Proceedingsof the Royal Society of London, Series B, 272, 1179–1183.
Rauhut, O. W. M. & Hungerbuhler, A. 2000. A review of Euro-pean Triassic theropods. GAIA, 15, 75–88.
Rauhut, O. W.M. & Lopez-Arbarello, A. 2009. Considerationson the age of the Tiouaren Formation (Iullemmeden Basin,Niger, Africa): implications for Gondwanan Mesozoic terres-trial vertebrate faunas. Palaeogeography, Palaeoclimatology,Palaeoecology, 271, 259–267.
Rauhut, O. W. M. & Werner, C. 1995. First record ofDromaeosauridae (Dinosauria: Theropoda) in the Creta-ceous of Gondwana (Wadi Milk Formation, northern Sudan).Palaontologische Zeitschrift, 69, 475–489.
Rauhut, O. W. M. & Xu, X. 2005. The small theropod dinosaursTugulusaurus and Phaedrolosaurus from the Early Creta-ceous of Xinjiang, China. Journal of Vertebrate Paleontology,25, 107–118.
Rauhut, O. W. M., Milner, A. C. & Moore-Fay, S. 2010. Cranialosteology and phylogenetic position of the theropod dinosaurProceratosaurus bradleyi (Woodward, 1910) from the MiddleJurassic of England. Zoological Journal of the Linnean Soci-ety, 158, 155–195.
Ray, G.E. 1941. Big for his day. Natural History, 41, 36–39.Reynolds, S. H. 1939. On a collection of reptilian bones from
the Oolite of Stow-on-the-Wold, Gloucestershire. GeologicalMagazine, 76, 193–214.
Riabinin, A. N. 1915. Zamtka o dinozavry ise Zabaykalya. TrudyGeologichyeskago Muszeyah Imeni Petra Velikago Impera-torskoy Academiy Nauk, 8(5), 133–140.
Riabinin, A. N. 1931. Pozvonki dinozavra iz nizhnego melaPrikaspiyskikh stepey. Zapiski Rossiyskogo Mineralogich-eskogo Obshchestva, 60, 110–113.
Rich, T. H. V. 1996. Significance of polar dinosaurs in Gondwana.Memoirs of the Queensland Museum, 39, 711–717.
Rich, T. H. V. & Rich, P.V. 1989. Polar dinosaurs and biotasof the Early Cretaceous of southeastern Australia. NationalGeographic Research, 5, 15–53.
Rich, T. H. V. & Vickers-Rich. 2003. Protoceratopsian? ulnaefrom Australia. Records of the Queen Victoria Museum, 113,1–12.
Rinehart, L. F., Lucas, S. G. & Hunt, A. P. 2007. Furculaein the Late Triassic theropod dinosaur Coelophysis bauri.Palaontologische Zeitschrift, 81, 174–180.
Rinehart, L. F., Lucas, S. G., Heckert, A. B., Spielmann, J. A. &Celeskey, M. D. 2009. The Paleobiology of Coelophysis bauri(Cope) from the Upper Triassic (Apachean) Whitaker Quarry,New Mexico, with Detailed Analysis of a Single Quarry Block.New Mexico Museum of Natural History and Science Bulletin,45, 1–260.
Ritgen, F. A. von 1826. Versuchte Herstellung einiger Beckenurweltlichter Thiere. Nova Acta Academiae CaesareaLeopoldino-Carolinae Germanicae Naturae Curiosorum, 13,331–358.
Romer, A. S. 1956. Osteology of the Reptiles. University ofChicago Press, Chicago, 772 pp.
Romer, A. S. 1966. Vertebrate Paleontology. University ofChicago Press, Chicago, 468 pp.
Rowe, T. B. & Gauthier, J. A. 1990. Ceratosauria. Pp. 151–168in D. B. Weishampel, P. Dodson & H. Osmolska (eds) TheDinosauria. University of California Press, Berkeley.
Ruiz-Omenaca, J. I. & Canudo, J. I. 2001. Dos yacimientosexcepcionales con vertebrados continentales del Barremiense(Cretacico Inferior) de Teruel: Vallipon y La Cantalera. Natu-raleza Aragonesa, 7, 8–18.
Ruiz-Omenaca, J. I., Canudo, J. I., Aurell, M., Badenas, B.,Barco, J. L., Cuenca-Bescos, G. & Ipas, J. 2004. Estado delas investigaciones sobre los vertebrados del Jurasico Superiory Cretacico Interior de Galve (Teruel). Estudios Geologicos,60, 179–202.
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
288 M. T. Carrano et al.
Ruiz-Omenaca, J. I., Canudo, J. I. & Infante, P. 2005. Pres-encia de un posible alosaurido (Dinosauria: Theropoda) enel Cretacico Inferior (Barremiense inferior) de La Maca 3(Galve, Teruel). Pp. 117–118 in E. Bernaldez, E. Mayoral &A. Guerreiro dos Santos (eds) XXI Jornadas de la SociedadEspanola de Paleontologıa. Gestion e Investigacion de laPaleontologıa en el Siglo XXI, Sevilla. Junta de Andalucıa,Consejerıa de Cultura, Sevilla.
Russell, D. A. 1984. A check list of the families and genera ofNorth American dinosaurs. Syllogeus, 53, 1–35.
Russell, D. A. 1996. Isolated dinosaur bones from the MiddleCretaceous of the Tafilalt, Morocco. Bulletin du MuseumNational d’Histoire Naturelle, Paris, 4e serie, Section C, 18,349–402.
Russell, D. A. & Dong, Z. 1994. The affinities of a new thero-pod from the Alxa Desert, Inner Mongolia, People’s Republicof China. Canadian Journal of Earth Sciences, 30, 2107–2127.
Sadleir, R., Barrett, P. M. & Powell, H. P. 2008. The anatomyand systematics of Eustreptospondylus oxoniensis, a thero-pod dinosaur from the Middle Jurassic of Oxfordshire,England. Monograph of the Palaeontographical Society,London, 160(627, for 2006), 1–82.
Sahni, A. & Bajpai, S. 1988. Cretaceous–Tertiary boundaryevents: the fossil vertebrate, palaeomagnetic and radiomet-ric evidence from peninsular India. Journal of the GeologicalSociety of India, 32, 382–396.
Sahni, A., Rana, R. S., Kumar, K. & Loyal, R. S. 1984. Newstratigraphic nomenclature for the intertrappean beds of theNagpur region. Indian Geoscience Journal, 5, 55–58.
Salgado, L., De la Cruz, R., Suarez, M., Fernandez, M.,Gasparini, Z., Palma-Heldt, S. & Fanning, M. 2008. FirstLate Jurassic dinosaur bones from Chile. Journal of VertebratePaleontology, 28, 529–534.
Sampson, S. D. & Witmer, L. M. 2007. Craniofacial anatomyof Majungasaurus crenatissimus (Theropoda: Abelisauridae)from the Late Cretaceous of Madagascar. Society of VertebratePaleontology Memoir, 8, 32–102.
Sampson, S. D., Krause, D. W., Dodson, P. & Forster, C.A. 1996. The premaxilla of Majungasaurus (Dinosauria:Theropoda), with implications for Gondwanan paleobiogeog-raphy. Journal of Vertebrate Paleontology, 16, 601–605.
Sampson, S. D., Witmer, L. M., Forster, C. A., Krause, D.W., O’Connor, P. M., Dodson, P. & Ravoavy, F. 1998.Predatory dinosaur remains from Madagascar: implicationsfor the Cretaceous biogeography of Gondwana. Science, 280,1048–1051.
Sampson, S. D., Carrano, M. T. & Forster, C. A. 2001. A bizarrepredatory dinosaur from the Late Cretaceous of Madagascar.Nature, 409, 504–506.
Sanz, J. L., Buscalioni, A. D., Moratalla, J. J., Frances, V.& Anton, M. 1990. Los reptiles Mesozoicos del registroespanol. Monografias del Museo Nacional de Ciencias Natu-rales, 2, 1–79.
Sauvage, H. E. 1882. Recherches sur les reptiles trouves dans leGault du Nord de l’est du Bassin de Paris. Memoires de laSociete Geologique de France, serie 3, 2, 1–41.
Sauvage, H. E. 1897–1898. Vertebres Fossiles du Portugal.Contributions a L’Etude des Poissons et des Reptiles du Juras-sique et Cretacique. Direction des Travaux Geologiques duPortugal, Lisbon, 46 pp.
Schrank, E. 2005. Dinoflagellate cysts and associatedaquatic palynomorphs from the Tendaguru Beds (UpperJurassic–Lower Cretaceous) of southeast Tanzania. Palynol-ogy, 29, 49–85.
Seeley, H. G. 1881. The reptile fauna of the Gosau Forma-tion preserved in the Geological Museum of the Universityof Vienna. Quarterly Journal of the Geological Society ofLondon, 37, 620–707.
Seeley, H. G. 1887. On Aristosuchus pusillus (Owen), beingfurther notes on the fossils described by Sir R. Owen as Poik-ilopleuron pusillus, Owen. Quarterly Journal of the Geologi-cal Society of London, 43, 221–228.
Sereno, P. C. 1999. The evolution of dinosaurs. Science, 284,2137–2147.
Sereno, P. C. 2007. The phylogenetic relationships of earlydinosaurs: a comparative report. Historical Biology, 19,145–155.
Sereno, P. C. & Brusatte, S. L. 2008. Basal abelisaurid andcarcharodontosaurid theropods from the Lower CretaceousElrhaz Formation of Niger. Acta Palaeontologica Polonica,53, 15–46.
Sereno, P. C., Forster, C. A., Rogers, R. R. & Monetta, A. M.1993. Primitive dinosaur skeleton from Argentina and theearly evolution of Dinosauria. Nature, 361, 64–66.
Sereno, P. C., Forster, C. A., Larsson, H. C. E., Dutheil, D. B.& Sues, H.-D. 1994. Early Cretaceous dinosaurs from theSahara. Science, 266, 267–271.
Sereno, P. C., Dutheil, D. B., Iarochene, M., Larsson, H. C.E., Lyon, G. H., Magwene, P. M., Sidor, C. A., Varricchio,D. J. & Wilson, J. A. 1996. Predatory dinosaurs from theSahara and Late Cretaceous faunal differentiation. Science,272, 986–991.
Sereno, P. C., Beck, A. L., Dutheil, D. B., Gado, B., Lars-son, H. C. E., Rauhut, O. W. M., Sadleir, R. W., Sidor,C. A., Varricchio, D. J., Wilson, G. P. & Wilson, J.A. 1998. A long-snouted predatory dinosaur from Africaand the evolution of spinosaurids. Science, 282, 1298–1302.
Sereno, P. C., Wilson, J. A. & Conrad, J. L. 2004. New dinosaurslink southern landmasses in the mid-Cretaceous. Proceedingsof the Royal Society of London, Series B, 271, 1325–1330.
Sereno, P. C., Martinez, R. N., Wilson, J. A., Varricchio, D. J.,Alcober, O. A. & Larsson, H. C. E. L. 2008. Evidence foravian intrathoracic air sacs in a new predatory dinosaur fromArgentina. PLoS ONE, 3(9), e3303, 1–20.
Siegwarth, J. D., Lindbeck, R. A., Redman, P. D., Southwell,E. H. & Bakker, R. T. 1997. Giant carnivorous dinosaurs ofthe family Megalosauridae from the Late Jurassic MorrisonFormation of eastern Wyoming. Contributions from the TateMuseum Collections, Casper, Wyoming, 2, 1–33.
Sikes, D. S. & Lewis, P. O. 2001. PAUPRat: PAUP∗ implemen-tation of the parsimony ratchet. beta software, version 1.Distributed by the authors. Department of Ecology and Evolu-tionary Biology, University of Connecticut, Storrs.
Smith, D. K. 1998. A morphometric analysis of Allosaurus. Jour-nal of Vertebrate Paleontology, 18, 126–142.
Smith, N. D., Makovicky, P. J., Pol, D., Hammer, W. R. &Currie, P. J. 2007. Osteology of Cryolophosaurus ellioti(Dinosauria: Theropoda) from the Early Jurassic of Antarc-tica and implications for early theropod evolution. ZoologicalJournal of the Linnean Society, 151, 377–421.
Smith, N. D., Makovicky, P. J., Agnolin., F., Ezcurra, M.,Pais, D. & Salisbury, S. 2008. A Megaraptor-like thero-pod (Dinosauria: Tetanurae) in Australia; support for faunalexchange across eastern and western Gondwana in the mid-Cretaceous. Proceedings of the Royal Society, Series B, 275,2085–2093.
Steel, R. 1970. Encyclopedia of Paleoherpetology. Part 14.Saurischia. Gustav Fischer Verlag, Portland, 87 pp.
Dow
nloa
ded
by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
The phylogeny of Tetanurae 289
Stovall, J. W. 1938. The Morrison of Oklahoma and its dinosaurs.Journal of Geology, 46, 583–600.
Stovall, J. W. & Langston, W., Jr 1950. Acrocanthosaurusatokensis, a new genus and species of Lower CretaceousTheropoda from Oklahoma. The American Midland Natu-ralist, 43, 696–728.
Stromer, E. 1915. Ergebnisse der Forschungsreisen Prof. E.Stromers in den Wusten Agyptens. II. Wirbeltier-Reste derBaharıje-Stufe (unterstes Cenoman). 3. Das Original desTheropoden Spinosaurus aegyptiacus nov. gen., nov. spec.Abhandlungen der Koniglich Bayerischen Akademie derWissenschaften, Mathematisch-physikalische Klasse Abhand-lung, 28, 1–31.
Stromer, E. 1931. Ergebnisse der Forschungsreisen Prof. E.Stromers in den Wusten Agyptens. II. Wirbeltier-Resteder Baharıjestufe (unterstes Cenoman). 10. Ein Skelett-Rest von Carcharodontosaurus nov. gen. Abhandlungen derBayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung, Neue Folge, 9, 1–23.
Stromer, E. 1934. Ergebnisse der Forschungsreisen Prof. E.Stromers in den Wusten Agyptens. II. Wirbeltierreste derBaharıje-Stufe (unterstes Cenoman). 13. Dinosauria. Abhand-lungen der Bayerischen Akademie der Wissenschaften,Mathematisch-naturwissenschaftliche Abteilung, Neue Folge,22, 1–79.
Sues, H.-D. & Taquet, P. 1979. A pachycephalosaurid dinosaurfrom Madagascar and a Laurasia-Gondwanaland connectionin the Cretaceous. Nature, 279, 633–635.
Sues, H.-D., Frey, E., Martill, D. M. & Scott, D. M. 2002.Irritator challengeri, a spinosaurid (Dinosauria: Theropoda)from the Lower Cretaceous of Brazil. Journal of VertebratePaleontology, 22, 535–547.
Sweetman, S. C. 2004. The first record of veociraptorinedinosaurs (Saurischia, Theropoda) from the Wealden (EarlyCretaceous, Barremian) of southern England. CretaceousResearch, 25, 353–364.
Swinton, W. E. 1955. Megalosaurus, the Oxford dinosaur.Advancement of Science, 12, 130–134.
Swinton, W. E. 1970. The Dinosaurs. Wiley-Interscience, NewYork, 331 pp.
Swofford, D. L. 2002. PAUP∗: Phylogenetic Analysis Using Parsi-mony (∗and Other Methods). Version 4.0b10. Sinauer Asso-ciates, Sunderland.
Taquet, P. 1976. Geologie et paleontologie du gisement de Gado-ufaoua (Aptien du Niger). Cahiers de Paleontologie. Editionsdu Centre National de la Recherche Scientifique, Paris, 191pp.
Taquet, P. & Russell, D. A. 1998. New data on spinosauriddinosaurs from the Early Cretaceous of the Sahara. ComptesRendus de l’Academie des Sciences a Paris, Sciences de laTerre et des Planetes, 327, 347–353.
Taquet, P. & Welles, S. P. 1977. Redescription du crane dedinosaure theropode de Dives (Normandie). Annales dePaleontologie (Vertebres), 63, 191–206.
Tawadros, E. 2001. Geology of Egypt and Libya. A. A. Balkema,Rotterdam, 468 pp.
Terquem, O. 1855. Paleontologie de l’etage inferieur de la Forma-tion Liasique de la Province de Luxembourg (Grand-Duche)et de Hettange, Departement de la Moselle. Memoires de laSociete Geologique de France, 5(2), 1–125.
Thevenin, Thevenin, A. 1907. Paleontologie de Madagascar.IV.—Dinosauriens. Annales de Paleontologie, 2, 121–136.
Torcida, F., Fuentes, C., Izquierdo, L. A., Montero, D. & Urien,V. 1997. Dientes de dinosaurios teropodos (cf. Baryonyx) en elWeald de Burgos (Espana). Stvdia Geologica Salmanticensia,33, 59–65.
Torcida Fernandez-Baldor, F., Canudo, J. I., Huerta, P.,Montero Huerta, D., Contreras, R., Perez Martınez, G. &Urien Montero, V. 2006. Un nuevo dinosaurio del CretacicoInferior de Burgos: el sauropodo de Oterillo II. Pp. 183–185in E. Fernandez-Martınez (ed.) XXII Jornadas de la SociedadEspanola de Paleontologıa, Leon.
Turner, A. H., Pol, D., Clarke, J. A., Erickson, G. M. & Norell,M. A. 2007. A basal dromaeosaurid and size evolution preced-ing avian flight. Science, 317, 1378–1381.
Turner, C. E. & Peterson, F. 1999. Biostratigraphy of dinosaursin the Upper Jurassic Morrison Formation of the WesternInterior, U.S.A. Pp. 77–114 in D. D. Gillette (ed.) VertebratePaleontology in Utah. Utah Geological Survey MiscellaneousPublication, 99–1.
Turner, C. E. & Peterson, F. 2004. Reconstruction of the UpperJurassic Morrison Formation extinct ecosystem—a synthesis.Sedimentary Geology, 167, 309–355.
Tykoski, R. S. 2005. Vertebrate paleontology in the Arizona Juras-sic. Mesa Southwest Museum Bulletin, 11, 72–93.
Tykoski, R. S. & Rowe, T. 2004. Ceratosauria. Pp. 47–70 in D. B.Weishampel, P. Dodson & H. Osmolska (eds) The Dinosauria,Second edition. University of California Press, Berkeley.
Tykoski, R. S., Forster, C. A., Rowe, T., Sampson, S. D. &Darlington, M. 2002. A furcula in the coelophysoid theropodSyntarsus. Journal of Vertebrate Paleontology, 22, 728–733.
Valenciennes, A. 1863. D’une espece de Chelonien fossile d’ungenre nouveau, trouve dans la craie du cap la Heve par M.Lennier, du Havre, et decrit par M. A. Valenciennes. CompteRendu des Seances de l’Academie des Sciences, 46, 317–322.
Veralli, C. & Calvo, J. O. 2003. New findings of carcharo-dontosaurid teeth on Futalognko quarry (Upper Turonian),north Barreales Lake, Neuquen, Argentina. XIX JornadasArgentinas de Paleontologıa de Vertebrados. Ameghiniana40(4, supplement), 74R.
Veralli, C. & Calvo, J. O. 2004. Dientes de teropodos carcharo-dontosauridos del Turoniano superior-Coniaciano inferior delNeuquen, Patagonia, Argentina. Ameghiniana, 41, 587–590.
Vianey-Liaud, M., Jain, S. L. & Sahni, A. 1987. Dinosaureggshells (Saurischia) from the Late Cretaceous Intertrappeanand Lameta Formations (Deccan, India). Journal of VertebratePaleontology, 7, 408–424.
Vickers-Rich, P., Rich, T. H., Lanus, D. R., Rich, L. S. V. &Vacca, R. 1999. “Big tooth” from the Early Cretaceous ofChubut province, Patagonia: a possible carcharodontosaurid.Pp. 85–88 in Y. Tomida, T. H. Rich, & P. A. Vickers-Rich (eds)Proceedings of the Second Gondwanan Dinosaur Symposium.National Science Museum Monographs, Tokyo, 15.
Viera, L. I. & Torres, J. A. 1995. Presencia de Baryonyx walkeri(Saurischia, Theropoda) en el Weald de La Rioja (Espana).Nota previa. Munibe, 47, 57–61.
Volkheimer, W., Quattrocchio, M., Cabaleri, N. & Garcıa, V.2008. Palynology and paleoenvironment of the Jurassic lacus-trine Canadon Asfalto Formation at Canadon Lahuinco local-ity, Chubut Province, central Patagonia, Argentina. RevistaEspanola de Micropaleontologıa, 40, 77–96.
Wagner, J. A. 1861. Neue Beitrage zur Kenntniss der urweltlichenFauna des lithographischen Schiefers. Abhandlungen derkoniglichen bayerische Akademie der Wissenschaft, II Classe,9, 65–124.
Waldman, M. 1974. Megalosaurids from the Bajocian (MiddleJurassic) of Dorset. Palaeontology, 17, 325–339.
Walker, A. D. 1964. Triassic reptiles from the Elgin area: Ornitho-suchus and the origin of carnosaurs. Philosophical Transac-tions of the Royal Society of London, Series B, 248, 53–134.
Wedel, M. J. 2003. Vertebral pneumaticity, air sacs, and the phys-iology of sauropod dinosaurs. Paleobiology, 29, 243–255.
Dow
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by [
Smith
soni
an I
nstit
utio
n L
ibra
ries
] at
07:
12 1
7 M
ay 2
012
290 M. T. Carrano et al.
Weishampel, D. B., Dodson, P. & Osmolska, H. 1990. TheDinosauria. University of California Press, Berkeley, 733 pp.
Weishampel, D. B., Dodson, P. & Osmolska, H. 2004a. TheDinosauria. Second edition. University of California Press,Berkeley, 861 pp.
Weishampel, D. B., Barrett, P. M., Coria, R. A., Le Loeuff,J., Xu, X., Zhao, X.-J., Sahni, A., Gomani, E. M. P. &Noto, C. R. 2004b. Dinosaur Distribution. Pp. 517–606 inD. B. Weishampel, P. Dodson & H. Osmolska, H. (eds) TheDinosauria. Second edition. University of California Press,Berkeley.
Welles, S. P. 1954. New Jurassic dinosaur from the KayentaFormation of Arizona. Bulletin of the Geological Society ofAmerica, 65, 591–598.
Welles, S. P. 1970. Dilophosaurus (Reptilia: Saurischia), a newname for a dinosaur. Journal of Paleontology, 44, 989.
Welles, S. P. 1983. Allosaurus (Saurischia, Theropoda) not yet inAustralia. Journal of Paleontology, 57, 196.
Welles, S. P. 1984. Dilophosaurus wetherilli (Dinosauria,Theropoda) osteology and comparisons. PalaeontographicaAbteilung A, 185, 85–180.
Wilkinson, M. 1995. Coping with abundant missing entries inphylogenetic inference using parsimony. Systematic Biology,44, 501–514.
Wilkinson, M. 2003. Missing data and multiple trees: stabilityand support. Journal of Vertebrate Paleontology, 23, 311–323.
Willison, S. W. 1878. American Jurassic dinosaurs. Transactionsof the Kansas Academy of Science, 6, 42–46.
Windolf, R. 1997. Theropoden-Zahne aus dem Oberen JuraNiedersachsens. Pp. 33–34 in S. Sachs, O. W. M. Rauhut &A. Weigert (eds) 1. Treffen der deutschsprachigen Palaoher-petologen, Terra Nostra, Extended Abstracts. Dusseldorf,Germany.
Wings, O., Broschinski, A. & Knotschke, N. 2005. New thero-pod and ornithopod dinosaur trackways from the Berriasianof Munchehagen (Lower Saxony, Germany). Kaupia, 14, 105.
Woodward, A. S. 1906. On a tooth of Ceratodus and a dinosaurianclaw from the Lower Jurassic of Victoria, Australia. Annalsand Magazine of Natural History, Decade 7, 18, 1–3.
Woodward, A. S. 1910. On a skull of Megalosaurus from theGreat Oolite of Minchinhampton (Gliucestershire). QuarterlyJournal of the Geological Society of London, 66, 111–115.
Woodward, A. S. & Sherborn, C. D. 1890. A Catalogue of BritishFossil Vertebrata. Dulao & Company, London, 396 pp.
Wu, X.-C., Currie, P. J., Dong, Z., Pan, S. & Wang, T. 2009. Anew theropod dinosaur from the Middle Jurassic of Lufeng,Yunnan, China. Acta Geologica Sinica, 83, 9–24.
Xu, X., Clark, J. M., Mo, J., Choiniere, J., Forster, C. A.,Erickson, G. M., Hone, D. W. E., Sullivan, C., Eberth, D.A., Nesbitt, S., Zhao, Q., Hernandez, R., Jia, C.-k., Han,F.-l. & Guo, Y. 2009. A Jurassic ceratosaur from China helpsclarify avian digital homologies. Nature, 459, 940–944.
Yates, A. M. 2005. A new theropod dinosaur from the Early Juras-sic of South Africa and its implications for the early evolutionof theropods. Palaeontologia Africana, 41, 105–122.
Young, C.-C. 1942. Fossil vertebrates from Kuangyuan, N.Szechuan, China. Bulletin of the Geological Society of China,22, 293–309.
Young, C.-C. 1958. [The first record of dinosaurian remains fromShansi]. Vertebrata PalAsiatica, 2, 231–236 [In Chinese].
Zhao, X.-J. 1986. [The Jurassic system of China]. Stratigraphy ofChina, 11, 286–348 [In Chinese].
Zhao, X.-J. & Currie, P. J. 1994. A large crested theropod fromthe Jurassic of Xinjiang, People’s Republic of China. Cana-dian Journal of Earth Sciences, 30, 2027–2036.
Zhao, X.-J., Benson, R. B. J., Brusatte, S. L. & Currie,P. J. 2010. The postcranial skeleton of Monolophosaurusjiangi (Dinosauria: Theropoda) from the Middle Jurassic ofXinjiang, China, and a review of Middle Jurassic Chinesetheropods. Geological Magazine, 147, 13–27.
Zittel, F. von. 1911. Grundzuge der Palaontologie (Palaozoolo-gie). II. Abteilung. Vertebrata. Druck und Verlag von R.Oldenbourg, Munchen, 598 pp.
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Appendix 1
Taxonomic hierarchy of Theropoda derived from the results of the present analyses. For each family-level taxon, the originalauthor(s) of the first associated family-level name is given immediately after the name, followed by the author(s) who firstcreated 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, 1986Coelophysoidea Holtz, 1994
Dilophosaurus wetherilli Welles, 1970Coelophysidae (Nopcsa, 1928) Paul, 1988
Coelophysis Cope, 1889C. bauri (Cope, 1887a) Cope, 1889C. rhodesiensis (Raath, 1969) Paul, 1988a
AVEROSTRA Paul, 2002 (sensu Ezcurra 2006)CERATOSAURIA Marsh, 1884
Elaphrosaurus bambergi Janensch, 1920Ceratosaurus nasicornis Marsh, 1884Abelisauroidea (Bonaparte & Novas, 1985) Bonaparte, 1991
Masiakasaurus knopfleri Sampson, Carrano & Forster, 2001Majungasaurus crenatissimus (Deperet, 1896) Lavocat, 1955
TETANURAE Gauthier, 1986 [syn. Avipoda Novas, 1992]‘Dilophosaurus’ sinensis Hu, 1993Cryolophosaurus ellioti Hammer & Hickerson, 1994
Monolophosaurus jiangi Zhao & Currie, 1994Chuandongocoelurus primitivus He, 1984ORIONIDES 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, 2005bMarshosaurus bicentesimus Madsen, 1976bPiatnitzkysaurus floresi Bonaparte, 1979
Megalosauria Bonaparte, 1850Streptospondylus altdorfensis Meyer, 1832 (Megalosauria incertae sedis)Spinosauridae Stromer, 1915
Baryonychinae (Charig & Milner, 1986) Sereno et al., 1998Baryonyx walkeri Charig & Milner, 1986Suchomimus tenerensis Sereno et al., 1998
Spinosaurinae (Stromer, 1915) Sereno et al., 1998Angaturama limai Kellner & Campos, 1996Irritator challengeri Martill et al., 1995Spinosaurus aegyptiacus Stromer, 1915
Megalosauridae Fitzinger, 1843Eustreptospondylus oxoniensis Walker, 1964Afrovenatorinae subfam. nov.
Afrovenator abakensis Sereno et al., 1994Dubreuillosaurus valesdunensis (Allain, 2002) Allain, 2005aLeshansaurus qianweiensis Li et al., 2009Magnosaurus nethercombensis (Huene, 1923) Huene, 1932Piveteausaurus divesensis (Walker, 1964) Taquet & Welles, 1977Poekilopleuron bucklandii Eudes-Deslongchamps, 1837
Megalosaurinae (Fitzinger, 1843) Paul, 1988Duriavenator hesperis (Waldman, 1974) Benson, 2008
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Megalosaurus bucklandii Mantell, 1827Torvosaurus 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, 1978Y. zigongensis (Gao, 1993) n. comb.
Metriacanthosaurinae (Paul, 1988a) subfam. nov.Metriacanthosaurus walkeri (Huene, 1923) Walker, 1964Shidaisaurus jinae Wu et al., 2009Siamotyrannus isanensis Buffetaut, Suteethorn & Tong, 1996Sinraptor Currie & Zhao, 1994
S. hepingensis (Gao, 1992) Currie & Zhao, 1994S. dongi Currie & Zhao, 1994
Allosauria Paul, 1988Allosauridae Marsh, 1878 [syn. Labrosauridae Marsh, 1882; Antrodemidae Stromer, 1934]
Allosaurus Marsh, 1877A. europaeus Mateus et al., 2006A. fragilis Marsh, 1877A. jimmadseni Chure et al., 2006
Saurophaganax maximus Chure, 1995Carcharodontosauria Benson et al., 2009
Neovenatoridae Benson et al., 2009Neovenator salerii Hutt, Martill & Barker, 1996Chilantaisaurus tashuikouensis Hu, 1964Megaraptora Benson et al., 2009
Aerosteon riocoloradensis Sereno et al., 2008Australovenator wintonensis Hocknull et al., 2009Fukuiraptor kitadaniensis Azuma & Currie, 2000Megaraptor namunhuaiquii Novas, 1998
Carcharodontosauridae Stromer, 1931Acrocanthosaurus atokensis Stovall & Langston, 1950Concavenator corcovatus Ortega, Escaso & Sanz, 2010Eocarcharia dinops Sereno & Brusatte, 2008Shaochilong maortuensis Brusatte et al., 2009Carcharodontosaurinae (Stromer, 1931) subfam. nov.
Carcharodontosaurus Stromer, 1931C. iguidensis Brusatte & Sereno, 2007C. saharicus (Deperet & Savornin, 1925) Stromer 1931
Giganotosaurus carolinii Coria & Salgado, 1995Mapusaurus roseae Coria & Currie, 2006Tyrannotitan chubutensis Novas et al., 2005
COELUROSAURIA Huene 1914Lourinhanosaurus antunesi Mateus, 1998Compsognathus longipes Wagner, 1861Ornitholestes hermanni Osborn, 1903Proceratosaurus bradleyi (Woodward, 1910) Huene, 1926
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Appendix 2
Taxon-by-character matrix for the 61 taxa used in this study. The two outgroup taxa are listed first, followed by the 59ingroup taxa listed alphabetically. Characters are scored from 0 to 1, 2, 3, or 4. Higher numbers represent uncodable characterobservations for particular taxa. Polymorphisms were assumed to represent uncertainties.
Eoraptor 00010 000?0 0000? ????? 01010 00?0? 00100 00?00 00100 ??000 1?000 0001000000 ?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?00000000 00000 0?000 0001? 00100 00?00 00010 00000 10100 00000 00?00 10000 0100000000 00001 00000 00?00 00000 0000? ?000? 00000 00?00 000?0 000?0 00?00 0000000??? 80000 00000 00000 00000 00000 0
Herrerasaurus 00001 00000 0000? ???0? 00010 00?0? 0?000 00?00 00100 ??001 2010000000 00000 ?0000 01000 00000 020?0 00010 ?0000 0?10? ?0000 0000? 00000 0?10000?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 1010001000 00000 00001 00000 00?00 00000 0000? 0?001 00000 0?000 00??0 00010 0000000000 001?? 90100 00100 00000 00000 00000 0
Acrocanthosaurus 020?1 100?0 00000 11110 00000 21111 11000 10?00 00002 11101 2111000202 12011 ?1121 11102 0110? 02??1 10121 00000 10011 01000 12011 111?1 1100210010 10200 10?11 111?0 00100 01000 00100 11012 ?0011 0011? 00?00 12110 0010001100 11211 110?0 02100 00001 01?1? 11011 01200 12001 210?0 01011 11000 0113110111 11100 0???? 010?1 0??11 10110 20120 1?120 ?2011 11000 22011 10111 1000011001 0312? 1010? 1110? 1??01 21011 1001? 1
Aerosteon ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?01?000?11 1???? ????? ?1000 0200? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? 11012 1???? ???11 ?0100 1?1?0 00100 0101101011 1???? ?2?11 0??0? ????? ???01 01100 121?? ????? ????? ????? ????? ?????????? 01011 01001 02211 10110 20120 1?120 ????? ????? ????? ????? ????? ?1001?412? ???01 11202 1110? ????? ????? ?
Afrovenator ????? ????0 01010 11000 00100 12001 1000? ????0 ?0002 01001 ?011001011 000?? ????? ??00? 00111 1001? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ??000 00000 01??? ??10? 11001 ?0?01 10000 10001 10110 0001?001?0 00001 ????? ??100 000?? ?1?01 010?? ????? ???01 21??? ??00? 0???? ?0?3100111 1?100 001?1 0?011 01011 ?011? 2012? 1?00? 00101 11000 11101 10000 1100001000 021?1 10101 10??? 11101 1?011 100?? 1
Allosaurus 00101 [01]0000 01000 10100 00000 21111 11000 11011 00002 [01]2[01]0010110 00000 01011 11100 01102 00000 02001 10110 00110 10010 01020 12011 1111111200 01001 10201 11111 11100 00100 01102 00100 11001 10011 0010[01] 10000 1011000100 00100 01011 11000 011?0 00000 0101? 11011 01200 12001 21010 11000 0100011131 00111 11100 00111 01001 02111 10110 20120 1?110 02001 11000 [12]1011 1011110000 11001 02122 10101 11102 11101 21011 10010 1
Angaturama 11??? 201?1 121?1 1???0 ????? ????? ???0? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????1 ?0???????? ????? ????? 11011 17013 11??? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ?
Australovenator ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
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?0000 1???? ????? ??100 00100 0110? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????0 ????? ????? ????? ????? ????? ???00 01010 1????01?11 11??1 01??? ????? ????? ????? ????? ????? ????? ????? 22012 10111 1000111001 14122 10101 11212 11101 211?? ??010 ?
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 101010?[12]10 1???1 ??010 01011 00013 00101 1100[12] ?1001 00100 10001 10111 1101000110 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 110100???? ????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 101111??01 ?100? ????? ?010? ????? ????? ????? ????? ?
Ceratosaurus 02101 00010 00000 00000 00000 01010 00101 20?00 00002 02001 1000000010 00011 11110 00101 00100 10001 01010 00000 00110 10000 11100 00?10 0010000001 ??100 00001 00100 01000 00101 00100 01102 1?010 00100 00000 02110 0000010100 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]001102100 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 1000100000 ?1000 041?2 12101 1???2 1110? 100?? 100?? ?
Coelophysis bauri 00101 21100 0010? ???0? 01101 0100? 00000 00?00 00101 ??00000000 00000 000?0 ?0000 01101 10100 000?0 00000 0?0?2 ??1?? ??0?? 1?0?0 ?0?00??001 10?0? 0000? 00?00 00000 01000 0?000 00000 01100 ?1000 11000 ?0000 0101100000 0?000 00000 01000 00000 0011? 1100? 01?00 00010 02000 10100 0000? 0[01]?0010111 01010 00000 10010 00110 00011 01111 0100? 10000 00100 00100 00110 [01]00?00?000 ?0100 01100 02100 00001 01111 00011 00000 0
Coelophysis rhodesiensis 00??? 21100 0010? ??00? 01101 100?0 00000 00?02 0010100001 [01]0000 0000? 00010 00000 00?01 10?00 00000 00000 00002 00100 ?000? 1000000000 10001 00000 00000 00000 00100 00000 01000 00000 01101 1?000 11000 0000001011 00000 00?00 00000 01000 0?000 0?11? 11000 01?00 00010 02000 10100 0?0010[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?000?00? 00??? ?1?00 ????? ???0? ?0??? ?0??0 0???? ?0??0 ?0??? ???1? 1???? 1?000
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0?00? ?0??1 00??1 0100? 0000? 0??00 00100 ?100[12] ????? ??100 ?001? ?0011 00?000???? 0000? ?1??0 ??000 ?0?00 ?1?0? 0111? 01110 0?0?? [12]??0? ??0?? ?0??0 1?1310011? ???0? 000?? 010?1 1??11 0011? 20?20 1?011 ?11?1 0??00 2???1 11??? ??????1?0? ????? ?0101 1???? ???01 ??0?1 1?010 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 010001???? ????? ????? 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 0001000?20 01?01 11010 01?00 02??? 01??1 01??? ????? ????? ????? ????? ????? ?????????? ????? 0?1?? 01?01 010?? ????? ?0??? ?0??? 00001 100?? ????? ?0??0 1000001?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 0100000000 ?2??? ???01 00101 1??0? ????? ????? ?
‘D. sinensis’ ?0?0? 000?0 010?? ????? 00020 2??0? 1??0? ?1013 00100 ??000 ?00?00?01? 000?? ?1??0 ????? 1?0?? ????? ????? ????? ????? ????? ????? ????? ????00??0? 00??? 1???? ????? 0??0? 0???? ??10? 0100[12] ?1?00 ??000 10000 ?0010 00000100?0 00001 ??000 ?1000 020?? 01?01 0??01 0001? 00011 21110 00000 01000 1???101011 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? 200100001? 00010 01000 ?1101 10100 00000 00?00 00002 00101 00000 1??0? ?0??? ??20110001 0000? 10000 00[01]10 00000 01000 00101 01101 ??000 11100 10000 01011 0000000000 00101 01000 01000 0201? ?1000 01000 00010 01001 10100 00001 01000 1?11101100 01000 10010 00110 00011 00110 [01]000? 10000 01000 1?001 00110 [01]000000000 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?? ????010?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? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????110101 ????? ????? ??000 0010? 0??0? 00100 ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ?
Elaphrosaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
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????? ????? ????? ????? ????? ????? ????? 0110[12] ????? ???00 10000 0?011 000001?000 00000 02111 ?0000 011?? 010?? ????? 0???1 1?011 10100 00100 0???? ?????????? ????? 00011 10000 00?11 11111 00?[12]? ????? 0?0?0 00011 00110 ?0011 0??0001000 0210? ?21?0 ????? ?111? 001?? ?1??? ?
Eocarcharia ????? ????0 00000 11100 10100 21111 11?0? ????? ????? ????? ??????01?0 02011 11??? 1???? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ??100 0010? 01??? ??10? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ?
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 1011000001 00110 00101 ?100? ?1?00 00??? ?1??? ????? 111?0 ???01 21210 ?00?? ?????????? ????? ????? ?0011 01011 00011 ?0113 00?2? 1?000 00101 0100? 11111 1000011000 01000 02111 101?1 10102 111?? 1001? ?000? ?
Fukuiraptor ????? ????? ????? ????0 ?0?0? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????0???? ????? ????? ??100 0010? 0110? ???00 110?[12] ????? ???1? ????? ????0 ????0????? ???11 ????? ????? ????? ????? ????? ????? 12001 21110 1?001 ?1010 ?????????? ????1 0???? ?10?? ????? ????? ????? ????? ????? ????? 21012 101?1 100011??01 ??12? ???01 11212 1??01 2?1?? ?0??? ?
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??201010 ????? 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]???? 100001???1 ?10?? ?2111 10?0? ????? ????? ????? ??0?? ?
Lourinhanosaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? 1100? ????? ???0? ????? 1?1?0 00?000?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
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10?11 ????? ????? ??000 0000? 0???? ????0 ???0? ????? ????? ????? ????? ?????????0 ????? ????? ???00 ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ?0??? 01??1 ????? ????? ????? ??0?? 0???? ????? 1???? ???0? 110?001000 ?21?? ???0? 11??? ????? ????? ????? ?
Majungasaurus 02101 00010 00000 10000 00010 01000 00111 20?10 02000 0?101 2000000202 02012 ?1111 01101 10000 10001 0100? 00000 010?0 1101? 10000 00010 1011001010 01100 10001 00101 00000 00100 00100 01102 10011 00100 11000 12110 0000010100 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?10001010 02102 11111 11102 1111? 0001? ?1101 ?
Mapusaurus ????? ?0??? 00000 11110 00010 01101 ?1010 11110 10002 1110? ?1111 102?212112 1???? ????[02] 01?00 0?0?? ????? ????? ????? ????? ????? ????? ??002 01010?0100 1???? ??100 00200 00?0? ??100 11012 ???01 ???1? ?0?00 10??0 001?? 01?0010201 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?1001?10 000?? ??100 0000? 10?0? 01?01 00020 00000 1001? 0?00? 1?0?? ????? ????10001? ?0??? 1?011 ??001 00100 01000 00100 11001 ?0111 11000 10000 12110 0000000120 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?? ????? ??11101000 01190 ??00? 00010 00000 00??? 00200 01102 ?0011 00100 11000 111?0 0000010100 00000 02011 11000 1011? 01000 01?01 01111 10011 10100 001?? ????? ?????????? ???00 1001? 10000 00011 ?1111 11120 10000 10010 01??1 00110 10011 0210001010 02102 ?1111 11202 1111? 0011? ?1101 ?
Megalosaurus ????? ????0 000?0 11011 00?00 110?1 ?000? ????? ????? ????? ?1?111???? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????000101 ????? 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?00001000 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?00011? 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?1111000 1??0? ??21? ????? ????? ????? ????? ????? ?
Monolophosaurus 001?1 000?0 0100? ????? 00000 2000? 10001 11013 00100 ?90012[01]110 00110 010?1 11100 01?02 10000 00??1 00010 00000 10??? ??0?? ??01? 1????
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??001 00001 00201 00?0? ??00? 0010? 0??00 00101 11001 0?001 00100 10?00 1011000000 0?100 00101 ?10?0 ?1?00 0?00? 010?? ????? ????? ????? ????? ????? ?????????? ????? ????? 00110 00101 01011 1011? 00?[12]0 100?? ?0001 000?0 ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ?
Neovenator 00??? 10000 01000 10101 00000 21001 ???10 11011 0???? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????1 ????? ????110000 ????? ????? ??100 00100 01?02 00100 11012 ?1??? ??111 10100 12110 0010001101 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 1000111001 13122 1010? 10??? 1???? 21011 1?010 ?
Ornitholestes 000?? 000?0 0000? ????? 00000 2100? 1?000 01000 00101 0?001 2?11000001 000?1 ??100 01?02 0?00? ?0??1 000?0 0???0 ????0 ??0?? ????? ????? ??00000?0? ?0?0? 0?010 0111? 00000 0??00 00100 1100[12] ????? ????0 10010 1?0?0 00?0001000 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 100001???? ???2? ?0?0? ????? ????? 2001? ?0??? ?
Piatnitzkysaurus ????? ????? 000?0 10000 10020 110?1 ???0? ????? ????? ????? ?????????? ????? ?1??? 0???? ????? ????1 0?010 00000 10?11 01000 ?10?? ????? ????2010?1 ????? ????? ??001 0010? 01??? ??000 01001 ?0011 11000 10000 12110 0001000120 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 1000001000 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?? ????? 201100000? ????? ????? ?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???011001 ?21?? ????? ????? ????? ?00?? ????? ?
Shaochilong ????? ????? 000?? 11?10 10010 01101 11?00 ???1? ????? ????? ?????????? ????? ?1?01 1???? ?0?00 02001 1???1 ?0000 100?0 011?? 120?? ????? ?????
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????? ????? ????? ??1?? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ?
Shidaisaurus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ???0? ?1100 0???? ????? ????1 10?0? ????? ????0 ????? ????? ????? ?????????? ????? ????? ????? 0??0? 00??? ????? ???0? ?0110 000?? ????? 1???? 001?00???? 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 0111000100 01001 ?1100 00112 00100 02001 10110 00100 10010 01120 10011 1?111 1100000011 00200 11111 00000 0010? 0?100 00100 11001 10110 00100 10000 12110 0011000110 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 1000011000 02212 10101 10102 11101 21011 10010 1
Sinraptor hepingensis 00001 00000 0000? ????? 01000 2111? 1?000 11011 00002 1100001110 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 1211000110 001?0 01211 ?1100 01100 0100? 0101? 11001 01110 0?0?? ????? ????? ?????????? ????? ????? 00001 01001 01?11 10112 [12]0021 11010 02101 10?10 110?1 101111?00? ????? ????? ????? ????? ????? ????? ????? ?
Spinosaurus 110?1 20101 1210? ???0? ?0000 020?? 1??0? ????0 ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ??00110?0? ?011? ????? ??010 11011 18013 11?10 1100[12] ????? ????? 1000? ?21?0 100?00???0 ?0201 0?00? ???00 0?0?? 0???? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ?
Streptospondylus ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? 1100[12] ????? ???0? ?0?01 ??1?0 000010?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 0011000201 ?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’ ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
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????? ????? ????? ????? 00?0? ?1??? ??00? 1100[12] ????? 0010? ?0000 12110 001000?1?0 0?011 ?1??0 ??000 ?100? ????? ???01 011?0 ???01 10210 01000 01000 1??2?00111 111?? 00001 010?1 0??11 10110 101[12]0 ?100? 02001 10111 11?1? ?0??1 10?000?00? ?211[12] 10?0? ?010? ????? ????? ????? ?
Torvosaurus 00??? 100?0 010?? 11011 00010 12000 ??00? ???00 ?1012 00001 ?011111011 000?[01] ????? ????? 00111 1?01? ????? ????? ????? ????? ????? ????? ????010?0? ????? ????? ??000 00100 01001 00100 11011 01021 00100 10001 12110 0001100110 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 1100001000 02121 10101 10102 1110? 1001? ?0??? ?
Tyrannotitan ????? ????? ????? ????? ????? ????? ???1? ????? ????? ????? ?1111????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????20?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?01???? ???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 011100010? 010?1 ?1??0 ?0?12 0???0 ?0??? ????? ????? ????? ????? ??011 1???1 ??00000?1? 00??? 1???1 ?0??? 00?0? 0??00 00100 1100[12] ?0110 001?0 10000 12110 001000?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???000[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 ?
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