Coelophysoidea Nopcsa, 1928 sensu Holtz, 1994
Definition- (Coelophysis bauri <- Carnotaurus sastrei) (modified from Sereno, 1998)
Other definitions- (Coelophysis bauri <- Ceratosaurus nasicornis) (modified from Padian et al., 1999)
(Coelophysis bauri <- Passer domesticus) (Allain et al., 2012)
(Coelophysis bauri <- Ceratosaurus nasicornis, Carnotaurus sastrei, Passer domesticus) (Sereno, in press)
= Podokesauroidea Huene, 1914 sensu Madsen and Welles, 2000
= Ceratosauria sensu Sereno, 1998
Definition- (Coelophysis bauri <- Passer domesticus) (modified)
= Coelophysoidea sensu Padian et al., 1999
Definition- (Coelophysis bauri <- Ceratosaurus nasicornis) (modified)
= Coelophysoidea sensu Sereno, in press
(Coelophysis bauri <- Ceratosaurus nasicornis, Carnotaurus sastrei, Passer domesticus)
Diagnosis- (after Tykoski, 2005) supracetabular crest flares lateroventrally to hide dorsal acetabulum in lateral view; anterior trochanter conical.
Comments- This clade may be part of Ceratosauria (Tykoski, 2005). See Comments under Ceratosauria.
History of use and Podokesauroidea- The idea of grouping Dilophosaurus with coelophysoids has a long history, but prior to 1994 was done under two older names- Podokesauridae (Russell, 1984; Carroll, 1988) and Coelophysidae (Paul, 1988; Novas, 1991; Novas, 1992). The clade was left unnamed by Rowe (1989), where it was first supported cladistically. Welles (1984) and some other authors had recognized the relationship between Dilophosaurus and Liliensternus, generally placing them both in Halticosauridae, but had only vague ideas about their relationship with smaller coelophysids. Holtz (1994) was the first to use the name Coelophysoidea for a Dilophosaurus + Coelophysidae clade, which was soon defined to be stem-based (Sereno, 1998) and thus included Coelophysis and any taxa more closely related to it than to neoceratosaurs. The term has been used in this way ever since. Madsen and Welles (2000) correctly noted that Podokesauroidea has priority over Coelophysoidea according to the ICZN, since it was named fourteen years earlier. Yet all other workers ignore this, generally because they value Phylocode rules and Podokesauroidea has not been phylogenetically defined, or they misread the ICZN to demand family-level eponyms be diagnosable (it actually only says they must be valid). Coelophysoidea is only used on this website because of its near universal usage since 1994.
Is Dilophosaurus a coelophysoid?- An additional phylogenetic issue is whether Dilophosaurus and related taxa (dilophosaurids as used here) are coelophysoids, or more closely related to ceratosaurs and tetanurines. Traditional phylogenies placed them in Coelophysoidea (see above), and this has been found in many recent analyses as well (Wilson et al., 2003; Holtz et al., 2004; Tykoski and Rowe, 2004; Tykoski, 2005; Ezcurra and Novas, 2006). Yet a few recent analyses have instead recovered Dilophosaurus as more closely related to ceratosaurs and tetanurines (Rauhut, 2003; Yates, 2005; Smith et al., 2007). Smith et al. recovered the following unambiguous synapomorphies for a dilophosaurid+ceratosaur+tetanurine clade- dorsoventrally elongate orbit; maxillae that are orientated subparallel to each other in dorsal aspect; reduced maxillary tooth count; tongue-like process of the parietals overlapping the supraoccipital knob; relatively deep surangular. Yet Rauhut found Dilophosaurus to be equally well supported as a coelophysoid when Shuvosaurus was (correctly) excluded, Yates found dilophosaurids were coelophysoids with only one added step, and Smith et al. found dilophosaurids were coelophysoids with only six added steps. Similarly, Ezcurra and Novas found equal bootstrap support for either option. Yet Tykoski (2005) found it took twenty additional steps to place Dilophosaurus closer to ceratosaurs and tetanurines. Perhaps importantly, none of the analyses supporting coelophysoid dilophosaurids have included Cryolophosaurus, Dracovenator or "Dilophosaurus" sinensis, whereas Smith et al. included all three and Yates includes Dracovenator. Also relevent is that Tykoski found Dilophosaurus to have been miscoded in prior analyses, due to a reliance on subadult specimens. Whether this was corrected for in later analyses has yet to be examined. If dilophosaurids are not coelophysoids, the coelophysoid identity of other taxa such as Sarcosaurus, Halticosaurus, Lophostropheus and Zupaysaurus comes into question as well. Indeed, both Smith et al. and Yates found Zupaysaurus to be outside of Coelophysoidea in their analyses.
Ex-coelophysoids- Several taxa have been included in authors' equivalents of Coelophysoidea in the past, but do not appear to belong there. Avipes, Lukousaurus, Saltopus and Velocipes were all included in Carroll's Podokesauridae, as was common in the pre-cladistic age. None have ever been placed in Coelophysoidea based on synapomorphies, the basic rationale being that they were small Triassic theropods. Current research suggests Saltopus is a more basal dinosauriform (e.g. Langer, 2004), and Avipes and Velocipes to be avemetatarsalians of more dubious nature (Rauhut and Hungerbuhler, 2000). Lukousaurus has not been recently restudied, but lacks several coelophysoid apomorphies and is more likely crurotarsan. Elaphrosaurus was assigned to Coelophysoidea (as Coelophysidae) by a couple authors (Paul, 1988; Novas, 1992) before it was realized to be a ceratosaur sensu lato (Holtz, 1994), which has been the result of every published cladistic analysis. Paul (1988) also believed spinosaurids were derived from coelophysids, but the description of more complete spinosaurid specimens has led to their classification as tetanurines (Sereno et al., 1996; Charig and Milner, 1997). Rauhut (2003) found Shuvosaurus to fall out as a coelophysoid in his analysis, but the description of Effigia (Nesbitt and Norell, 2006) led to its recognition as a crurotarsan.
Coelophysoidea defined- Sereno's (1998) and Padian et al.'s (1999) definitions for Coelophysodea are basically equivalent as nearly every phylogeny agrees Ceratosaurus and Carnotaurus are more closely related to each other than either is to Coelophysis. The presence of Passer as a specifier in Sereno's (in press) redefinition is useful, but Carnotaurus seems superfluous, since there has never been a (Ceratosaurus (Passer (Carnotaurus, Coelophysis))) topology suggested. Allain et al. (2012) suggest only Passer as the external specifier, which leads to difficulties is coelophysoids are ceratosaurs, as then Coelophysoidea is a synonym of Ceratosauria instead of a subgroup.
References- Huene, 1914. Beiträge zur geschichte der Archosaurier [Contribution to the history of the archosaurs]. Geologie und Paläontologie Abhandlungen. 13(7), 1-56
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Russell, 1984. A check list of the families and genera of North American dinosaurs. Syllogeus. 53, 1-35.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Carroll, 1988. Vertebrate Paleontology and Evolution. W.H. Freeman and Company.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. J. Vert. Paleontol. 9, 125-136.
Novas, 1991. Relaciones filogeneticas de los dinosaurios teropodos ceratosaurios [Phylogenetic relationships of ceratosaurian theropod dinosaurs]. 28(3-4), 401.
Novas, 1992. La evolucion de los dinosaurios carnivoros [The evolution of carnivorous dinosaurs]. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico: Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez", Cuenca, Argentina. 126-163.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Charig and Milner, 1997. Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum of London (Geology). 53, 11-70.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. N. Jb. Geol. Paläont. Abh. 210(1), 41-83.
Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998. A long-snouted predatory dinosaur from Africa and the evolution of the spinosaurids. Science. 282(5392), 1298-1302.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature of the major taxonomic categories of the carnivorous Dinosauria (Theropoda). Journal of Vertebrate Paleontology, 19(1), 69-80.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised osteology. Miscellaneous Publication 00-2, Utah Geological Survey. 80 pp.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia. 15, 75-88.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid (Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian) of India. Contr. Mus. Palaeont. Univ. Mich. 31, 1-42.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods. Palaeontologia Africana. 41, 105-122.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article, 38 pp. DOI 10.1080/08912960600845791.
Nesbitt and Norell, 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B. 273, 1045-1048.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften. 99(5), 369-377.

unnamed possible coelophysoid (Galton, 1976)
Hettangian, Early Jurassic
Shuttle Meadow Formation, Connecticut, US
Material- tooth
Comments- Galton (1976) referred this to Coelophysis sp., but Coelophysis teeth are not known to be diagnostic.
Reference- Galton, 1976. Prosauropod dinosaurs (Reptilia: Saurischia) of North America. Postilla. 169:1-98.

unnamed coelophysoid (Kirby, 1991)
Rhaetian, Late Triassic
Owl Rock Member of the Chinle Formation, Arizona, US
Material- (MNA V7240) proximal femur
Comments- This was assigned to Ceratosauria? by Kirby (1991) and was labeled Coelophysis sp. in the MNA collections. Hunt et al. (1998) mentioned it as undoubtedly dinosaurian, and Spielmann et al. (2007) used the hooked femoral head and strong trochanteric shelf to refer it to Coelophysoidea.
References- Kirby, 1991. A vertebrate fauna from the Upper Triassic Owl Rock Member of the Chinle Formation of northern Arizona. Unpublished Masters thesis. Northern Arizona University, Flagstaff. 496 pp.
Kirby, 1993. Relationships of Late Triassic basin evolution and faunal replacement in the southwestern United States: Perspectives from the upper part of the Chinle Formation in northern Arizona. In Lucas and Morales (eds.). The Nonmarine Triassic. New Mexico Museum of Natural History and Science Bulletin. 3, 233-242.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the Western United States. Geobios. 31(4), 511-531.
Spielmann, Lucas and Heckert, 2007. Tetrapod fauna of the Upper Triassic (Revueltian) Owl Rock Formation, Chinle Group, Arizona. In Lucas and Spielmann (eds.). The Global Triassic, New Mexico Museum of Natural History and Science Bulletin. 41, 371-383.

undescribed possible coelophysoid (Ford, DML 1998)
Early Jurassic
Hanson Formation, Antarctica
Material- teeth
Comments- Although described as 'halticosaurid' by Ford, coelophysoid teeth are not known to be diagnostic.
Reference- http://dml.cmnh.org/1998Aug/msg00810.html

unnamed coelophysoid (Cunningham, Hungerbuhler, Chatterjee and McQuilkin, 2002)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US
Material- (TTUP coll.) tibia
Comments- Cunningham et al. (2002) identified this specimen as an ornithischian, but Nesbitt et al. (2007) noted it has theropod synapomorphies like a fibular crest, and shares a subrectangular distal outline with a small posterolateral process with coelophysoids. It is indistinguishable from Coelophysis and Gojirasaurus.
References- Cunningham, Hungerbuhler, Chatterjee and McQuilkin, 2002. Late Triassic vertebrates from the Patricia site near Post, Texas. Journal of Vertebrate Paleontology. 22(supplement to 3). 47A.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

unnamed possible coelophysoid (Maisch and Matzke, 2003)
Bathonian-Callovian, Middle Jurassic
Toutunhe Formation, Xinjiang, China
Material- (SGP 2000/2) (~3 m) distal fibula
(SGP 2001/5) (~3 m) tooth
Comments- Maisch et al. (2001) labeled the fibula as a coelurosaur. Maisch and Matzke (2003) note the fibula is most similar to Sarcosaurus and Megapnosaurus, based on the anteroproximal inclination of the distal articular facet. The tooth was assigned to Coelophysoidea based on the large number of serrations (33-37/mm), which is comparable to Coelophysis and Liliensternus.
Reference- Maisch and Matzke, 2003. Theropods (Dinosauria, Saurischia) from the Middle Jurassic Toutunhe Formation of the southern Junggar Basin, NW China. Palaeontologische Zeitschrift. 77(2):281-292.

Sarcosaurus Andrews, 1921
?= "Liassaurus" Welles, Powell and Pickering vide Pickering, 1995
S. woodi Andrews, 1921
= Magnosaurus woodi (Andrews, 1921) Huene, 1932
?= "Liassaurus huenei" Welles, Powell and Pickering vide Pickering, 1995
Sinemurian, Early Jurassic
Lower Lias, England
Holotype- (BMNH R4840/1) (robust) (~3.6 m, ~60 kg) partial posterior dorsal vertebra, partial ilia (248 mm), proximal pubes, ischial fragment, incomplete femur (~335 mm)
Referred- ?(Warwick Museum coll.) (gracile) (~3.5 m, ~55 g) anterior dorsal centrum (45 mm), mid dorsal centrum (50 mm), ilial fragment, partial pubes, incomplete femora (~320 mm), tibiae (one proximal; 297 mm), distal fibula, distal metatarsal II, distal metatarsal III, proximal phalanx III-1, distal metatarsal IV (Huene, 1932)
Diagnosis- (after Andrews, 1921) subpreacetabular notch extremely acute.
(proposed) dorsal ilial edge strongly convex; brevis fossa hidden in lateral view; postacetabular process convex posteriorly; medial femoral condyle small in posterior view; tibia posteriorly convex in distal view;
Other diagnoses- Andrews (1921) also distinguished Sarcosaurus from Ceratosaurus, Megalosaurus and Allosaurus by its small conical anterior trochanter, but this is plesiomorphic.
Paul (1988) stated the more proximally placed "outer ridge" (= trochanteric shelf?) on the femur distinguished it from Ceratosaurus, but this does not appear to be true.
Comments- Carrano and Sampson (2004) claimed Sarcosaurus is undiagnostic, but the ilium appears to possess apomorphies.
Huene (1932) assigned the species to Magnosaurus, though this is certainly incorrect as the latter is a tetanurine.
"Liassaurus"- In 1927, Huene was informed of a theropod partial skeleton in the Warwick Museum which he later described (1932) as a specimen of Sarcosaurus woodi. A distal pubis referred to the specimen is not theropod, as the end is extremely flattened anteroposteriorly. It is very similar to sauropodomorphs more derived than Saturnalia, but less than Yunnanosaurus, and is here referred to that clade. Carrano and Sampson (2004) referred to the specimen as cf. Sarcosaurus woodi, seemingly depending on Huene's description. While both Huene and Carrano and Sampson noted similarity between the Warwick specimen and the woodi holotype, neither provided synapomorphies to support such a referral. Pickering (1995) listed the name Liassaurus huenei in an unpublished bibliographic work, credited to Welles, Powell and Pickering. This is a nomen nudum however, as he didn't follow ICZN Article 8.1.3- it must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies. He later (Welles and Pickering, 1999) referred to it as an unnamed Liassic theropod in the comparative section of an unpublished Megalosaurus redescription. It is presumably one of the theropods to be redescribed by Welles and Powell in their unpublished study from the 1970's, which Pickering intends to publish as Mutanda Dinosaurologica. Pickering posted his diagnosis for "Liassaurus" on a private newsgroup in 2005, of which only the absent trochanteric shelf was different from Sarcosaurus. Comparing the two is difficult as only an ilial fragment, proximal pubes and incomplete femur are preserved in both. The ilial fragment is undescribed, and neither the pubis or femur preserve diagnostic characters or combinations of characters in both since both lack the femoral head, the woodi type lacks distal condyles and the Warwick specimen lacks the anterior trochanter. Characters which differ between femora (curvature, fourth trochanter lateromedial placement, presence of trochanteric shelf) vary within other basal theropod taxa as well. The sharp mediodistal flare on the tibia is similar to Elaphrosaurus and Ceratosaurus, while the distal tibial depth is greater than Megapnosaurus, ceratosaurs or tetanurines.
Relationships- Andrews (1921) originally assigned this genus to the Megalosauridae, in which he included not only basal tetanurines, but also Ceratosaurus and carnosaurs. Specifically, Andrews felt the ilium was similar to Ceratosaurus and Megalosaurus, while the anterior trochanter was more primitive than Allosaurus, Megalosaurus and Ceratosaurus, though closest to the latter. An assignment to such a broad Megalosauridae was standard throughout the 1900's. Kurzanov (1989) referred it to his more restricted concept of Megalosauridae (containing Megalosaurus and sinraptorids) without comment. Huene (1956) referred it to his incorrectly formed family Coelurosauridae. Gauthier (1986) recognized Sarcosaurus as a ceratosaur sensu lato on the basis of its trochanteric shelf, which was followed by Rowe (1989). This was elaborated on by Rowe and Gauthier (1990), who placed Sarcosaurus in Ceratosauria but outside their unnamed equivalent of Coelophysidae based on the poorly defined M. iliofemoralis fossa on the ilium and the lack of an 'obturator ridge' on the posterior femoral head. Paul (1988) referred Sarcosaurus to the Ceratosauridae based on pelvic bones which are fused and "nearly identical" in shape to Ceratosaurus. Welles (1984) was the first to suggest coelophysoid relations, placing it in Coelophysidae with Coelophysis (but not other currently recognized coelophysoids) without comment. Carrano and Sampson (2004) described characters shared with coelophysoids, such as long dorsal centra, an anteriorly facing pubic peduncle, small, spike-like anterior trochanter, and notched, circular distal tibia. They also noted resemblence to Liliensternus and Dilophosaurus, and referred Sarcosaurus to Coelophysoidea. Although Carrano et al. (2005) could not recover any resolution within Coelophysoidea in their cladistic analysis, when several characters are changed to ordered, the results differ. If a standard coelophysoid topology of (Dilophosaurus (Liliensternus (Coelophysis, Megapnosaurus))) is enforced, Sarcosaurus is placed outside the Liliensternus + Coelophysis + Megapnosaurus clade. Tykoski (2005) notes that Sarcosaurus has a mix of coelophysoid and ceratosaur sensu stricto characters, and found it to be the most basal coelophysoid in their analysis based on two characters- supracetabular crest flares lateroventrally to hide dorsal acetabulum in lateral view; anterior trochanter conical.
References- Andrews, 1921. On some remains of a theropodous dinosaur from the Lower Lias of Barrow-on-Soar. Annual Magazine of Natural History. 9(8), 570-576.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Huene, 1956. Paläontologie und Phylogenie der Niederen Tetrapoden. VEB Gustav Fischer Verlang, Jena. 1-716.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica Abteilung A. 185, 85-180.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Kurzanov, 1989. O proiskhozhdenii i evolyutsii infraotryada dinozavrov Carnosauria [Concerning the origin and evolution of the dinosaur infraorder Carnosauria]. Paleontologicheskiy Zhurnal. 1989(4), 3-14.
Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9, 125-136.
Rowe and Gauthier, 1990. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 151-168.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry. A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola, California. 478 pp.
Welles and Pickering, 1999. Megalosaurus bucklandii. Private publication of Stephen Pickering, An extract from Archosauromorpha: Cladistics & Osteologies. A Fractal Scaling in Dinosaurology Project. 119 pp.
Carrano and Sampson, 2004. A review of coelophysoids (Dinosauria: Theropoda) from the Early Jurassic of Europe, with comments on the late history of the Coelophysoidea. Neues Jahrbuch fur Geologie und Palaontologie Monatshefte. 2004, 537-558.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate Paleontology. 25(4), 835-849.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.

Gojirasaurus Carpenter, 1997
= "Revueltoraptor" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
G. quayi Carpenter, 1997
= "Revueltoraptor lucasi" Hunt 1994 vide Nesbitt, Irmis and Parker, 2007
Norian, Late Triassic
Bull Canyon (=Cooper Canyon) Formation of the Dockum Group, New Mexico, US
Holotype- (UCM 47221; in part) (5.5 m) pubis (497 mm), tibia (469 mm)
Diagnosis- Provisionally indeterminate relative to Coelophysis bauri.
Comments- This specimen was originally briefly described and illustrated as Procompsognathidae gen. et sp. indet. by Parrish and Carpenter (1986). It was then described and named "Revueltoraptor lucasi" by Hunt (1994) in his unpublished thesis, and called herrerasaurid A by Hunt et al. (1998). It was finally officially named and described as the coelophysoid Gojirasaurus quayi by Carpenter (1997). However, Nesbitt et al. (2005) find the holotype to be chimaeric, which was elaborated on in their 2007 publication. They found the dorsal vertebrae were referrable to Shuvosaurus and the pubis and tibia referrable to a coelophysoid (contra the 2005 abstract where the dinosaurian elements were relegated to Saurischia indet.). They could not assign the other material (teeth, dorsal ribs, gastralia, chevron) or the several referred specimens noted by Hunt (1994) (NMMNH P4666, P16607, P16656, P16946, P17134, P17154, P17258, UMMP 7274) to a particular taxon of archosaur. The reassignment of the dorsal material makes the sole apomorphy identified by Rauhut invalid (mid/posterior dorsal vertebrae with taller neural spines than other coelophysoids), and Gojirasaurus is presently indeterminate relative to Coelophysis except for the greater robustness of the tibia (Nesbitt et al., 2007). This may be size-related.
Although the name "Revueltoraptor lucasi" was originally used in thesis, and thus not available for use in this website, it was later published by Nesbitt et al. (2007). The name was first noted publically on the Dinosaur Mailing List in 2000 by Gay, who noted several elements at the NMMNH were labeled as "Revueltoraptor". These were specimens referred to the taxon by Hunt (1994), and are not identifiable as Gojirasaurus.
Rauhut (2003) suggested Gojirasaurus may be synonymous with Shuvosaurus, as a large Shuvosaurus premaxilla was found in the same deposits (identified as Reptilia indet. by Parrish and Carpenter, 1986), and the tooth associated with the Gojirasaurus holotype may not belong to it. However, Nesbitt and Norell (2006) have recently demonstrated that Shuvosaurus is a crurotarsan whose pubis and tibia differ markedly from Gojirasaurus'.
Relationships- Hunt (1994) and Hunt et al. (1998) referred this specimen to Herrerasauridae based on the short dorsal centra, strap-like scapula and elongate pubis. However, the dorsals are now assigned to Shuvosaurus, the scapula is Archosauria indet., and all theropods have elongate pubes. Parrish and Carpenter (1986) believed the taxon to be most closely related to Liliensternus within their Procompsognathidae, in which they included all coelophysoids. Carpenter (1997) concurred regarding the similarity to Liliensternus, referring it to Coelophysoidea and possibly Coelophysidae. Rauhut's analysis (2003) found Gojirasaurus to be a coelophysoid more closely related to coelophysids than to Liliensternus. Yates' (2005) updated version of Rauhut's analysis found Gojirasaurus to be in a polytomy with Megapnosaurus, "M." kayentakatae and Segisaurus, more derived than Coelophysis. Tykoski and Rowe (2004) could only say that Gojirasaurus was a coelophysoid more derived than Dilophosaurus, which was also the result of Carrano et al.'s (2005) analysis. Tykoski (2005) could find even less resolution, with Gojirasaurus having an uncertain position within Coelophysoidea, though in a polytomy with Zupaysaurus, Liliensternus and Coelophysis in the majority rule tree. Finally, Ezcurra and Novas (2006) found it to be the sister taxon of "Megapnosaurus" kayentakatae, outside Coelophysinae. However, all of these analyses included Shuvosaurus and Archosauria indet. remains in their Gojirasaurus OTUs, as they were published prior to Nesbitt et al.'s reanalysis of the holotype. Thus any particular placement within Coelophysoidea is questionable, and must be reexamined to determine if it was based on pubic or tibial characters.
References- Parrish and Carpenter, 1986. A new vertebrate fauna from the Dockum Formation (Late Triassic) of eastern New Mexico. In: Padian, K. (ed.). The Beginning of the Age of Dinosaurs. Cambridge Univ. Press, NewYork. Pp.151-160.
Hunt, 1994. Vertebrate paleontology and biostratigraphy of the Bull Canyon Formation (Chinle Group: Norian), east-central New Mexico with revisions of the families Metoposauridae (Amphibia: Temnospondyli) and Parasuchidae (Reptilia: Archosauria). Unpublished PhD Dissertation. Albuquerque, Univerrsity of New Mexico. 403 pp.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Carpenter, 1997. A giant coelophysoid (Ceratosauria) theropod from the Upper Triassic of New Mexico, USA. Neues Jahrbuch fuer Geologie und Palaeontologie, Abhandlungen. 205(2), 189-208.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.
http://dml.cmnh.org/2000Oct/msg00495.html
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate Paleontology. 25(4), 835–849.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. JVP 25(3) 96A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods. Palaeontologia Africana. 41, 105-122.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article, 38 pp. DOI 10.1080/08912960600845791.
Nesbitt and Norell, 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B. 273, 1045-1048.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

Halticosauridae Bock, 1952
Halticosaurinae Bock, 1952 sensu Paul, 1988
Comments- Bock (1952) originally named this as a family of coelurosaurs (sensu Huene) containing Liliensternus (as Halticosaurus liliensterni). It is often ascribed to Huene (1956). Welles (1984) included Dilophosaurus, Halticosaurus, Liliensternus and Longosaurus, while Chatterjee (1993) and Hu (1993) also included the first two genera. Paul (1988) used a subfamily Halticosaurinae to include Liliensternus and Dilophosaurus. The family has never been defined with explicit synapomorphies, instead being a receptacle for the larger coelophysoids which current analyses indicate form a grade basal to coelophysids. Although Halticosaurus has never been entered in a data matrix, Liliensternus and Dilophosaurus have never formed a clade exclusive of Coelophysis in any published study. Thus Halticosauridae has remained unused by most current workers. If Halticosaurus is shown in the future to share synapomorphies with Liliensternus, Dilophosaurus, Sarcosaurus or another taxon outside Coelophysidae, Halticosauridae should be applied to the resulting clade.
References- Bock, 1952. Triassic reptilian tracks and trends of locomotive evolution. Journal of Paleontology. 26(3), 395-433.
Huene, 1956. Paläontologie und Phylogenie der Niederen Tetrapoden [Paleontology and Phylogeny of the Lower Tetrapods]. VEB Gustav Fischer Verlang, Jena. 1-716.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Chatterjee, 1993. Shuvosaurus, a new theropod: an unusual theropod dinosaur from the Triassic of Texas. National Geographic Research and Exploration. 9(3), 274-285.
Hu, 1993. A new Theropoda (Dilophosaurus sinensis sp. nov.) from Yunnan, China. Vertebrata PalAsiatica. 31(1), 65-69.
Halticosaurus Huene, 1907
H. longotarsus Huene, 1907
Middle Norian, Late Triassic
Middle Stubensandstein, Germany
Holotype- (SMNS 12353) dentary fragment (lost), teeth (lost), sixth (?) cervical vertebra (57 mm), two cervical vertebrae (lost), anterior dorsal vertebra (33 mm) (lost), dorsal centrum (43 mm), second sacral vertebra (35 mm) (lost), third sacral vertebra (33 mm) (lost), proximal caudal vertebra (36 mm), proximal humerus (lost), ilial fragments (lost), proximal femora (300+ mm), metatarsal II (134 mm)
Referred- (HMN coll.) fragmentary remains (Huene, 1921)
Diagnosis- (after Welles, 1984) differs from Liliensternus liliensterni in- shallower dentary; dentary more pointed anteriorly; shorter and taller mid cervical vertebrae; sharp ventral keel on mid cervical vertebrae; narrower sacral centra; lower third sacral vertebra; more distally placed anterior trochanter; smaller distal condyles on metatarsal II which extend less far proximally on shaft.
Comments- Norman (1990) and Rauhut and Hungerbuhler (2000) believe Halticosaurus is indeterminate. However, Welles (1984) finds differences between comparable taxa such as Liliensternus and Dilophosaurus.
Relationships- Huene (1908) originally assigned Halticosaurus only to Dinosauria, then later (1909) to Saurischia. Romer's (1956) assignment to Hallopodidae is incorrect, as Hallopus is a crurotarsan differing from Halticosaurus and other theropods in having a proximally unprojected anterior trochanter among other characters. Norman (1990) believed the remains could be from a basal sauropodomorph or a theropod, though the sauropodomorph characters seems to be untrue (cervical pleurocoels absent; two sacral vertebrae; similar femur), and the theropod character primitive (elongate metatarsal). Assignments to Compsognathidae (Zittel, 1911) and Coeluridae (Huene, 1920) are similarly incorrect, as the low spike-like anterior trochanter is more primitive than coelurosaurs. Most workers have assigned Halticosaurus to Podokesauridae (Huene, 1914), an equivalent Procompsognathidae (Romer, 1966), or its eponymous family Halticosauridae (Bock, 1952). These categories are equivalent to the modern concepts of Coelophysoidea, with halticosaurids now thought to be a basal grade of that clade. Between 1934 and 1984, such statements were usually based on Liliensternus, then believed to be a species of Halticosaurus. Rauhut and Hungerbuhler (2000) give the only modern account of H. longotarsus, noting the material is very poorly preserved and most is not identifiable as theropod. As the holotype was found with Sellosaurus gracilis remains, some may be prosauropod. The proximal femora show a spike-like lesser trochanter and downturned head, as in coelophysoids. They therefore think some of the type may be coelophysoid. If this is true, the short cervical centrum excludes it from Coelophysidae. However, Halticosaurus has never been included in a phylogenetic analysis or described in detail since its discovery.
References- Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs of the European Triassic formations with consideration of occurrences outside Europe]. Geologische und Palaeontologische Abhandlungen Suppl. 1(1), 1-419.
Huene, 1909. Skizze zu einer Systematik und Stammesgeschichte der Dinosaurier [Sketch of the systematics and origins of the dinosaurs]. Centralblatt für Mineralogie, Geologie und Paläontologie. 1909, 12-22.
Zittel, 1911. Grundzüge der Paläontologie (Paläozoologie). II. Abteilung. Vertebrata [Fundamentals of Paleontology (Paleozoology). Section II. Vertebrata]. Druck und Verlag von R. Oldenbourg, München. 1-598.
Huene, 1914. Das natürliche System der Saurischia [The systematics of the Saurischia]. Centralblatt für Mineralogie, Geologie und Paläontologie. 1914, 154-158.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien [Phylogenetic results of some investigations of Triassic reptiles]. Zeitschrift für Induktive Abstammungsund Vererbungslehre. 24, 159-163.
Huene, 1921. Coelurosaurier-Reste aus dem obersten Keuper von Halberstadt. Cbl. Min. Geol. Palaeontol. 1921(10), 315-320.
Bock, 1952. Triassic reptilian tracks and trends of locomotive evolution. Journal of Paleontology. 26(3), 395-433.
Huene, 1956. Paläontologie und Phylogenie der Niederen Tetrapoden [Paleontology and Phylogeny of the Lower Tetrapods]. VEB Gustav Fischer Verlang, Jena. 1-716.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press, Chicago. 1-468.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie 185 p. 85-180.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. Univ. Cal. press. Berkeley.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia. 15, 75-88.

Sarcosaurus? andrewsi Huene, 1932
= Magnosaurus woodwardi Huene, 1932
= Megalosaurus (Magnosaurus) woodwardi Huene, 1932
= Megalosaurus andrewsi (Huene, 1932) Waldman, 1974
Sinemurian, Early Jurassic
Lower Lias, England
Holotype- (BMNH R3542) tibia (445 mm)
Diagnosis- Provisionally indeterminate relative to Sarcosaurus woodi.
Comments- Carrano and Sampson (2004) found it to be indistinguishable from the referred specimen of Sarcosaurus woodi except for size, and suggested the two species might be synonyms. The specimen was accidentally made the type of two species simultaneously by Huene (1932), who later (1956) used S. andrewsi for it. In his 1932 paper, Huene also lists Megalosaurus woodwardi in the section on Magnosaurus nethercombensis, stating both should be Megalosaurus subgenus Magnosaurus. As Magnosaurus is explicitly named as a new genus on that same page, this was probably an earlier opinion that was mistakenly retained.
References- Woodward, 1908. Note on a megalosaurian tibia from the Lower Lias of Wilmcote, Warwickshire. Annals and Magazine of Natural History. 8(1), 257-259.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monog. Geol. Pal. 4 (1) pts. 1 and 2, viii + 361 pp.
Huene, 1956. Palaeontologie und Phylogenie der niederen Tetrapoden. Fischer-Verlag, Jena.
Carrano and Sampson, 2004. A review of coelophysoids (Dinosauria: Theropoda) from the Early Jurassic of Europe, with comments on the late history of the Coelophysoidea. N. Jb. Geol. Palaont. Mh. 2004 (9): 537-558.

unnamed clade (Dilophosaurus wetherilli + Coelophysis bauri)
Diagnosis- (after Tykoski, 2005) skull elongate (>3 times longer than tall); antorbital fenestra elongate (>25% skull length); premaxillary tooth row ends anterior to external naris; elongate posteromedial process on premaxilla; premaxilla and maxilla loosely articulated; premaxillary and maxillary alveolar margins do not contact; anterior tip of maxillary alveolar margin curves mediodorsally; first maxillary alveolus opens anteroventrally; anteromedial maxillary process elongate; dorsal edge of anterior dentary raised; most premaxillary teeth lack serrations; anterior dentary teeth enlarged; narrow cervical epipophyses which project posterolaterally; no axial diapophyses; axial parapophyses reduced or absent; pubic peduncle of ilium with two facets- facing anteriorly and ventrally; distal tips of pubes lack contact; nasal process of premaxilla comprises >50% of external naris' dorsal edge; frontals elongated and triangular; lateral lamina of lacrimal sinuous and extends anteriorly past medial lamina; anterior pleurocoels in cervical vertebrae; elongate anterior dorsal centra (>1.33 height of anterior articular surface); elongate posterior dorsal centra (>1.33 height of anterior articular surface); ventral surface of caudal centra with narrow, sharp longitudinal groove; posterior scapular margin nearly straight over majority of length; distal carpals I and II fused; phalanx I-1 elongated (> 100% metacarpal I length); manual digit II longer than digit III; ischial antitrochanter large, protrudes anteriorly into acetabulum; obturator notch in ischium; oblique ridge on proximomedial fibula that overlaps groove.

Dilophosauridae Madsen and Welles, 2000
= "Dilophosauridae" Charig and Milner, 1990
= Dilophosaurinae Madsen and Welles, 2000
Diagnosis- (after Smith et al., 2007) contribution of the premaxillary posterodorsal process to a blade-like nasal crest; extension of the antorbital fossa onto the
lateroventral side of the nasal; presence of a nasolacrimal crest.
Comments- Although historically various taxa (Liliensternus, Halticosaurus, etc.) have been proposed to be more closely related to Dilophosaurus than to Coelophysis or tetanurines (e.g. Welles, 1984; Paul, 1988), these were not supported by phylogenetic analyses. The first analysis to recover such sister taxa to Dilophosaurus was that of Yates (2005), which found a clade containing Dilophosaurus, Zupaysaurus and Dracovenator to be sister to Ceratosauria+Tetanurae. Smith et al. (2007) later found a clade containing "Dilophosaurus" sinensis, Dracovenator, Cryolophosaurus and Dilophosaurus (but not Zupaysaurus, which was slightly more basal). Zupaysaurus has been assigned to Coelophysidae in other recent analyses (Carrano et al., 2005; Tykoski, 2005; Ezcurra and Novas, 2006). Smith et al.'s dilophosaur clade was also sister to Ceratosauria+Tetanurae. Both Yates' and Smith et al.'s analyses could place their dilophosaur clades in Coelophysoidea with few added steps (1 in Yates; 6 in Smith et al.).
Dilophosauridae was first used by Charig and Milner (1990) to refer to Paul's (1988) informal dilophosaurs, which he actually used the subfamiliy Halticosaurinae for (contra their quote). Since they did not diagnose or define the family, it is a nomen nudum (ICZN Article 13.1.1). Madsen and Welles (2000) mention a Dilophosauridae used in the same sense as Coelophysoidea. However, they then state Dilophosaurinae should be a subfamily of Podokesauridae containing only Dilophosaurus. Finally, they show an indented table with Dilophosauridae containing only Dilophosaurinae, with Dilophosaurus its sole genus. This table lists characters of Dilophosaurinae/idae, so this is the first valid use of the family. While neither Yates nor Smith et al. used a name for their dilophosaur clade, Dilophosauridae seems appropriate and is used here.
References- Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Charig and Milner, 1990. The systematic position of Baryonyx walkeri, in the light of Gauthier's reclassification of the Theropoda. in Carpenter and Currie (eds.). Dinosaur Systematics: Approaches and Perspectives. Cambridge University Press, Cambridge. 127-140.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised osteology. Miscellaneous Publication 00-2 Utah Geological Survey, 80 pages.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate Paleontology. 25(4), 835–849.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods. Palaeontologia Africana. 41, 105-122.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article, 38 pp. DOI 10.1080/08912960600845791.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.

"Dilophosaurus" sinensis Hu, 1993
= Dilophosaurus “sinensis” Dong, Hisa and Azuma, 1990
Hettangian, Early Jurassic
Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
Holotype- (KMV 8701) (5.5 m) skull (525 mm), lower jaw (487 mm), nine cervical vertebrae, cervical ribs, fifteen dorsal vertebrae, four sacral vertebrae, thirty-six caudal vertebrae, chevrons, scapulae, coracoids, distal clavicle, humerus, radius, ulna, metacarpals, ilium, pubis, ischium, femur (587 mm), tibia, fibula, astragalus, calcaneum, distal tarsal IV, metatarsus, pes
Referred- (LDM-LCA 10) skull, incomplete skeleton (Dong, 2003)
(ZLJ0003) partial skull, incomplete skeleton (Xing, 2012)
(ZLJT01) (immature) premaxillae fragment, incomplete maxilla, maxilla fragment, lacrimal, frontals, parietals, incomplete braincase, incomplete dentary, atlantal intercentrum, two dorsal rib fragments, partial proximal caudal neural arch (Xing, 2012)
Diagnosis- (after Carrano et al., 2012) vertical groove on lateral premaxilla adjacent to contact with maxilla.
(suggested) five premaxillary teeth.
Comments- The holotype was discovered in 1987, and has been confused with the popular mounted specimen LDM-LCA 10 which has a longer skull among other differences. Carrano et al. (2012) believe the differences are ontogenetic or taphonomic, as the skulls share the the diagnostic premaxillary groove, a vertical anterior border of the maxilla, and a similarly shaped and positioned promaxillary fenestra. Currie et al. (in prep.) ascribe the differences to sexual dimorphism, but this has yet to be published.
Lamanna et al. (1998) doubt this species is referrable to Dilophosaurus based on differences in premaxillary shape, tooth count, lack of antorbital tooth row, shape of infratemporal fenestra and squamosal and size and position of external mandibular fenestra. Dong (2003) believes this taxon is similar to, and perhaps synonymous with Sinosaurus triassicus, though without citing supporting evidence. Xing (2012) also synonymizes the taxa without evidence, and cites Currie et al. (in prep.) as supporting the synonymy.
Smith et al. (2007) included it in their phylogenetic analysis, where it emerged as the most basal dilophosaurid. Carrano et al. (2012) recovered it as a non-orionidan tetanurine, but only three more step remove it from Neotheropoda. Ten steps were needed to place it in Coelophysoidea where they recovered Dilophosaurus however. As several other relevent taxa were not included (e.g. Zupaysaurus, Dracovenator, Sarcosaurus), this result is questionable.
References- Hu, 1993. A new Theropoda (Dilophosaurus sinensis sp. nov.) from Yunnan, China. Vertebrata PalAsiatica. 31(1), 65-69.
Lamanna, Smith, You, Holtz and Dodson, 1998. A reassessment of the Chinese theropod dinosaur Dilophosaurus sinensis. Journal of Vertebrate Paleontology. 18(3), 57A.
Dong, 2003. Contributions of new dinosaur materials from China to dinosaurology. Memoir of the Fukui Prefectural Dinosaur Museum. 2, 123-131.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Xing, 2012. Sinosaurus from Southwestern China. Masters Thesis. University of Alberta. 267 pp.
Currie, Xing, Wu and Dong, in prep. Anatomy and relationships of Sinosaurus triassicus (“Dilophosaurus sinensis”) from the Lufeng Formation (Lower Jurassic) of Yunnan, China.

Sinosaurus Young, 1948
S. triassicus Young, 1948
Sinemurian, Early Jurassic
Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Holotype- (IVPP V34) two maxillary fragments, four maxillary teeth, jaw fragment, three teeth
Paratypes- (IVPP V35, V36, V37, V48) teeth
Hettangian, Early Jurassic
Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
Paratypes- ?(IVPP V30) incomplete mid dorsal vertebra (153 mm)
?(IVPP V31) partial posterior dorsal vertebra (~160 mm)
(IVPP V38) teeth
Hettangian-Sinemurian, Early Jurassic
Lufeng Formation, Yunnan, China
Referred- (CUP 2001) tooth (Simmons, 1965)
(CUP 2002) tooth (Simmons, 1965)
(CUP 2003) dorsal centrum (Simmons, 1965)
(CUP 2004) tooth (Simmons, 1965)
(CUP 2005) tooth (Simmons, 1965)
(CUP 2095) dorsal centrum (Simmons, 1965)
(CUP 2096) three tooth fragments (Simmons, 1965)
(CUP 2097) anterior jaw fragment, teeth (Simmons, 1965)
(CUP 2098) cervical centrum, two dorsal centra (Simmons, 1965)
Comments- Dong (2003) believes this taxon is similar to, and perhaps synonymous with "Dilophosaurus" sinensis, though without citing supporting evidence. Xing (2012) also synonymizes the taxa without evidence, and cites Currie et al. (in prep.) as supporting the synonymy.
The postcrania referred to the taxon by Young (1948; 1951) are sauropodomorph (Walker, 1964), later assigned to Jingshanosaurus (Zhang and Yang, 1995), but the latter referral was doubted by Galton and Upchurch (2004).
Carrano et al. (2012) recovered sinensis as a non-orionidan tetanurine, but only three more steps remove it from Neotheropoda. Ten steps were needed to place it in Coelophysoidea where they recovered Dilophosaurus however. As several other relevent taxa were not included (e.g. Zupaysaurus, Dracovenator, Sarcosaurus), this result is questionable.
References- Young, 1948. On two new saurischians from Lufeng, Yunnan. Bulletin of the Geological Society of China. 28 75-90.
Young, 1951. The Lufeng saurischian fauna in China. Palaeontologica Sinica. C(13), 1-96.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London B. 248, 53-134.
Simmons, 1965. The non-therapsid reptiles of the Lufeng Basin, Yunnan, China. Fieldiana Geology. 15, 1-93.
Zhang and Yang, 1995. A complete osteology of Prosauropoda in Lufeng Basin Yunnan China. Jingshanosaurus. Yunnan Science and Technology Publishing House, Kunming. 100 pp.
Dong, 2003. Contributions of new dinosaur materials from China to dinosaurology. Memoir of the Fukui Prefectural Dinosaur Museum. 2, 123-131.
Galton and Upchurch, 2004. Prosauropoda. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Xing, 2012. Sinosaurus from Southwestern China. Masters Thesis. University of Alberta. 267 pp.
Currie, Xing, Wu and Dong, in prep. Anatomy and relationships of Sinosaurus triassicus (“Dilophosaurus sinensis”) from the Lufeng Formation (Lower Jurassic) of Yunnan, China.

unnamed clade (Dracovenator regenti + Cryolophosaurus ellioti + Dilophosaurus wetherilli)
Diagnosis- (after Smith et al., 2007) low height/length ratio of the premaxilla below the external naris; slot-shaped foramen on the lateral face of the premaxilla at the base of the nasal process (unknown in "Dilophosaurus" sinensis); presence of erect, tab-like dorsal processes on the articular (unknown in "Dilophosaurus" sinensis).

Dracovenator Yates, 2005
D. regenti Yates, 2005
Hettangian-Sinemurian, Early Jurassic
Upper Elliot Formation, South Africa
Holotype- (BP/1/5243) premaxillae, maxillary fragment, two dentary fragments, partial surangular, partial angular, partial prearticular, articular, teeth
Paratype- ?(BP/1/5278) (juvenile) premaxillae, incomplete maxillae, partial nasals, partial dentaries, teeth (Munyikwa and Raath, 1999)
Diagnosis- (after Yates, 2005) a large bilobed fossa surrounding a large lateral premaxillary foramen that is connected to the alveolar margin by a deep narrow channel; a deep, oblique notch on the lateral surface of the articular, separating the retroarticular process from the posterior margin of the glenoid; particularly well-developed dorsal, tab-like processes on the articular, one on the medial side, just posterior to the opening of the chorda tympanic foramen and the other on the lateral side on the anterolateral margin of the fossa for the m. depressor mandibulae.
References- Munyikwa and Raath, 1999. Further material of the ceratosaurian dinosaur Syntarsus from the Elliot Formation (Early Jurassic) of South Africa. Palaeontologia Africana. 35:55-59.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods. Palaeontologia Africana. 41, 105-122.

unnamed clade (Cryolophosaurus ellioti + Dilophosaurus wetherilli)
Diagnosis- (after Smith et al., 2007) lateral groove along the posterior end of the surangular.

Cryolophosaurus Hammer and Hickerson, 1994
= "Elvisaurus" Holmes, 1993
C. ellioti Hammer and Hickerson, 1994
Rhaetian-Toarcian, Late Triassic-Early Jurassic
Hanson Formation, Antarctica
Holotype- (FMNH PR1821) (~6.5 m; 465 kg) (partial skull ~460 mm) two maxillary fragments, nine maxillary teeth, posterior skull, posterior mandibles, sixth cervical central fragment, seventh cervical vertebra (118 mm), eighth cervical vertebra (108 mm), ninth cervical vertebra, tenth cervical vertebra, several posterior cervical ribs, several anterior dorsal vertebrae, most mid and posterior dorsal vertebrae (114, 125, 115, 117 mm), several dorsal ribs, fifth sacral vertebra (89 mm), partial first caudal centrum, two mid caudal vertebrae (100 mm), three mid caudal centra, many partial and complete caudal vertebrae, three chevrons, partial humeri, proximal radius, proximal ulna, partial ilium, proximal pubis, ischia (one distal), incomplete femora (769 mm), distal tibia, distal fibula, astragalus, calcaneum
Diagnosis- (after Smith et al., 2007) large, anterodorsally curving midline crest with fluted rostral and caudal surfaces formed by dorsal expansions of the lacrimals; complete constriction across the infratemporal fenestra formed by the squamosal and jugal; extremely elongate cranial processes on the cervical ribs.
Comments- Smith et al. (2007) include a note added in proof that the supposed proximal tibia they describe is a proximal humerus.
Smith et al. (2005) concluded this was a very basal tetanurine, but their later published analysis (Smith et al., 2007) found Cryolophosaurus to clade with Dilophosaurus, "D." sinensis and Dracovenator in a Dilophosauridae. Carrano et al. (2012) have since recovered it as a non-orionidan tetanurine again, but only one more step removes it from Neotheropoda. Nine steps were needed to place it in Coelophysoidea where they recovered Dilophosaurus however. As several other relevent taxa were not included (e.g. Zupaysaurus, Dracovenator, Sarcosaurus), this result is questionable.
References- Holmes, 1993. Prehistoric Times.
Hammer and Hickerson, 1994. A crested theropod dinosaur from Antarctica. Science. 264, 828-830.
Smith, Hammer and Currie, 2005. Osteology and phylogenetic relationships of Cryolophosaurus ellioti (Dinosauria: Theropoda): Implications for basal theropod evolution. Journal of Vertebrate Paleontology. 25(3), 116A-117A.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

Dilophosaurus Welles, 1970
D. wetherilli (Welles, 1954) Welles, 1970
= Megalosaurus wetherilli Welles, 1954
= Dilophosaurus "breedorum" Welles, 1995 vide Welles and Pickering, 1995
Hettangian, Early Jurassic
Silty Facies Member of Kayenta Formation, Arizona, US
Holotype- (UCMP 37302) (young subadult) (6.03 m, 283 kg) skull (523 mm), atlas (19 mm), axial intercentrum (17 mm), axis (56 mm), third cervical vertebra (~75 mm), fourth cervical vertebra (80 mm), fifth cervical neural arch, sixth cervical vertebra (86 mm), seventh cervical neural arch, eighth cervical neural arch (~87 mm), posterior ninth cervical centrum (~88 mm), posterior tenth cervical centrum (~80 mm), first dorsal centrum (73 mm), second dorsal vertebra (67 mm), third dorsal vertebra (~62 mm), fourth dorsal vertebra (67 mm), fifth dorsal vertebra (74 mm), sixth dorsal vertebra (70 mm), seventh dorsal vertebra (78 mm), eighth dorsal vertebra (84 mm), ninth dorsal vertebra (82 mm), tenth dorsal vertebra (~88 mm), eleventh dorsal vertebra (86 mm), twelfth dorsal vertebra (90 mm), thirteenth dorsal vertebra (70 mm), dorsal ribs, partial first sacral vertebra, second sacral vertebra (69 mm), third sacral vertebra (67 mm), fourth sacral vertebra (67 mm), fifth sacral vertebra (70 mm), first caudal vertebra (69 mm), second caudal vertebra (65 mm), third caudal vertebra (65 mm), caudals 4-44, chevrons 2-36, scapulae (375 mm), coracoids (105 mm high, 185 mm long), humeri (285, 270 mm), radii (180, 192 mm), ulnae (205, 209 mm), radiale, distal carpal I, distal carpal II, distal carpal III, distal carpal IV, carpal, metacarpal I (51 mm), phalanx I-1 (69 mm), manual ungual I (70 mm), metacarpal II (105 mm), phalanx II-1 (70 mm), phalanx II-2 (63 mm), manual ungual II (47 mm), metacarpal III (115 mm), phalanx III-1 (41 mm), phalanx III-2 (44 mm), phalanx III-3 (45 mm), manual ungual III (38 mm), metacarpal IV (68 mm), phalanx IV-1 (21 mm), metacarpal V, ilia (370 mm), pubes (485 mm), ischia (340 mm), femur (557 mm), tibia (555 mm), fibula (518 mm), astragalus (92 mm wide, 77 mm tall), calcaneum, two distal tarsals, metatarsal I (95 mm), phalanx I-I (68 mm), pedal ungual I (55 mm), metatarsal II (250 mm), phalanx II-1 (103 mm), phalanx II-2 (79 mm), pedal ungual II (73 mm), metatarsal III (300 mm), phalanx III-1 (110 mm), phalanx III-2 (84 mm), phalanx III-3 (70 mm), pedal ungual III (70 mm), metatarsal IV (254 mm), phalanx IV-1 (72 mm), phalanx IV-2 (57 mm), phalanx IV-3 (46 mm), phalanx IV-4 (38 mm), pedal ungual IV (45 mm), metatarsal V (126 mm)
Paratypes- (UCMP 37303) (young subadult) partial skull (550 mm) including premaxillae, maxillae, palatine, ectopterygoid, basioccipital, dentary, splenial, prearticular and teeth, partial fifth cervical vertebra, sixth cervical vertebra (92 mm), ninth dorsal vertebra (87 mm), tenth dorsal vertebra (92 mm), first caudal vertebra (70 mm), second caudal vertebra (73 mm), third caudal centrum (77 mm), metacarpal I (48 mm), partial phalanx I-1, metacarpal II (90 mm), incomplete metacarpal III, metacarpal III (88 mm), partial metacarpal IV, partial distal tibia, distal fibula, astragalus, calcaneum
Referred- (MNA P1.97) tooth (Gay, 2001)
(MNA P1.101) proximal pubis (Gay, 2001)
(MNA P1.102) (juvenile) distal fibula (Gay, 2001)
(MNA P1.109) (juvenile) femur (281 mm) (Gay, 2001)
(MNA P1.111) dentary fragments (Gay, 2001)
(MNA P1.116) manual phalanx III-1 (48 mm) (Gay, 2001)
(MNA P1.122) ischial fragment (Gay, 2001)
(MNA P1.131) pedal phalanx III-1 (120 mm) (Gay, 2001)
(MNA P1.135) dorsal centrum (83 mm), partial neural spine (Gay, 2001)
(MNA P1.138) dorsal centrum (45 mm) (Gay, 2001)
(MNA P1.141) manual phalanx III-2 (60 mm) (Gay, 2001)
(MNA P1.142) partial rib (Gay, 2001)
(MNA P1.154) proximal pubis (Gay, 2001)
(MNA P1.160) proximal femur (~545 mm) (Gay, 2001)
....(MNA P1.161) distal femur (Gay, 2001)
(MNA P1.176) dorsal centrum (31 mm) (Gay, 2001)
(MNA P1.177) dorsal centrum (85 mm), partial neural spine (Gay, 2001)
(MNA P1.247) partial rib (Gay, 2001)
(MNA P1.248) pubis (570 mm) (Gay, 2001)
(MNA P1.530) distal fibula (Gay, 2001)
(MNA P1.539) distal fibula (Gay, 2001)
(MNA P1.3145) proximal femur (~574 mm) (Gay, 2005)
(MNA P1.3181) (juvenile) tooth fragment, distal humerus, distal fibula (Gay, 2001)
?(TMM 43646) (~5.1 m; young subadult) partial skeleton including partial maxilla, partial braincase, fourth cervical vertebra, dorsal neural arch, five sacral neural arches, proximal caudal vertebrae, mid and distal caudal vertebrae, pectoral girdle, ilia, incomplete pubis, ischia, hindlimbs including femora (443 mm), tibia, fibula, astragalus, calcaneum, distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, metatarsal III and metatarsal IV (Tykoski, 2005)
(TMM 43662-2) (gracile adult) proximal femur (Tykoski, 2005)
(UCMP 77270; intended holotype of Dilophosaurus "breedorum") (~6.4 m; ~340 kg; robust adult) incomplete skull (619 mm), mandible (635 mm), anterior dentary, atlantal centrum (17 mm) plus axis (88 mm), incomplete third cervical vertebra (77 mm), fourth cervical vertebra (80 mm), fifth cervical vertebra (90 mm), sixth cervical vertebra (104 mm), seventh cervical vertebra (100 mm), eighth cervical vertebra, ninth cervical vertebra (82 mm), tenth cervical vertebra (92 mm), atlantal rib (295 mm), axial rib, seven cervical ribs, incomplete second dorsal vertebra (75 mm), partial third dorsal vertebra (86 mm), partial fourth dorsal vertebra (87 mm), fragmentary fifth dorsal vertebra, fragmentary sixth dorsal vertebra, partial seventh dorsal vertebra, partial eighth dorsal vertebra, partial ninth dorsal vertebra, partial tenth dorsal vertebra (97 mm), eleventh dorsal vertebra (92 mm), incomplete thirteenth dorsal vertebra (83 mm), second dorsal rib (370 mm), fifth dorsal rib (450 mm), sixth dorsal rib, seventh dorsal rib, tenth dorsal rib (280 mm), eleventh dorsal rib (280 mm), twelfth dorsal rib fragment, gastralial fragments, partial sacrum (78, 86, ?, ?, ? mm), sixth or seventh caudal centrum (87 mm), few caudal vertebrae, scapulocoracoid, radius (180 mm), incomplete ulna (215 mm), partial ilium fused to proximal pubis, distal pubis, fragmentary ischia, femur (590 mm), tibia (558 mm), proximal fibula, incomplete metatarsal I (~106 mm), metatarsal II (248 mm), pedal ungual II (70 mm), metatarsal III (300 mm), pedal ungual III (68 mm), metatarsal V (125 mm) (Welles, 1970)
(UCMP 130053) incomplete skeleton including vertebral fragments (Clark and Fastovsky, 1986)
(UCMP 130069) ischium (UCMP online)
(UCMP 130070) vertebrae, proximal femur, limb fragment, phalanx (UCMP online)
(UCMP 130083) scapulocoracoid fragments (UCMP online)
Diagnosis- (after Rauhut, 2000) lacrimal with thickened dorso-posterior rim; cervical neural spines with a distinct central "cap" and an anterior and posterior "shoulder"; scapular blade with squared distal expansion.
(after Carrano et al., 2012) thin, paired nasolacrimal crests extending vertically from skull roof, each with fingerlike posterior projection.
Comments- The Paleobiology Database is responsible for the specimen number and identification of UCMP 130053, otherwise mentioned an an undescribed theropod in Clark and Fastovsky (1986).
Xu et al. (2009) illustrated metacarpal V in the holotype. Based on Welles' (1984) description and the morphology in Coelophysis, his radiale is distal carpal I, ulnare is distal carpal II, intermedium is the radiale, 'second carpale' is distal carpal III, one of the other ossicles is distal carpal IV, while the other may be an intermedium.
Carrano et al. (2012) note that TMM 43646 differs from D. wetherilli in a few characters (taller maxillary interdental places; pneumatic fossa on the dorsal surface of the jugal process of the maxilla), and excluded it from the hypodigm in their analysis.
Dilophosaurus "breedorum"- The incomplete skeleton UCMP 77270 was discovered in 1964 which was initially mentioned by Welles (1970) as a larger specimen of Dilophosaurus wetherilli. This was the first specimen of the genus to preserve a nearly complete cranial crest, the bases of which had been preserved but unnoticed in the holotype. Welles later (1984) believed UCMP 77270 to be a new related genus of theropod based on undescribed differences in skull proportions, vertebrae and especially the femur. Gauthier (1986) retained the specimen in D. wetherilli and stated Welles had remarked on its trochanteric shelf in 1984, yet as Charig and Milner (1990) noted, Gauthier was mistaken and Welles never described the femur of the specimen. Rowe and Gauthier (1990) also referred it to D. wetherilli and incorrectly stated it was of similar ontogenetic stage as the holotype. Paul (1988) was not certain whether it was the same species or not, but noted the differences might be due to the same kind of dimorphism that coelophysids show. Welles wrote a description of UCMP 77270 in which he names it Dilophosaurus breedorum, which was eventually released by Pickering in 1995. This paper has controversial status, as it describes the only one of Pickering's taxa to be accepted as valid by another paleontologist (Olshevsky, DML online 1999). Olshevsky noted that the publication had no evidence of following ICZN Articles 8.1.2 and 8.1.3, but considered D. "breedorum" valid on the condition that Pickering could supply copies in response to orders. While I have received a copy from Pickering, he has refused to send them to several other workers or to archive them in public libraries. Thus Olshevsky's condition has been only partially met, and whether he still considers the species to be valid is unknown. Other workers such as Ford (Paleofile.com) consider "breedorum" a nomen nudum. Incidentally, Olshevsky used the date 1999 for the "breedorum" paper and stated 1995 "must be a manuscript date, since the description was not published then and has only appeared through Pickering's efforts this year (1999)." I personally do not doubt Pickering printed the "breedorum" paper in 1995, though its distribution at that time is questionable. I provisionally accept the 1995 date here, though I also consider the taxon a nomen nudum under Article 8.1. Note that contra Olshevsky, if the "breedorum" paper is accepted as valid under the ICZN, "Newtonsaurus" and "Walkersaurus" from the comparative section would also be valid. In any case, "breedorum" was definitely used in Pickering's 1995 bibliographic work "Jurassic Park: Unauthorized Jewish Fractals in Philopatry" as a nomen nudum. In that work, the name is a label for a photograph of UCMP 77270's skull. Gay (2005) believed it was a specimen of D. wetherilli, feeling there is a "lack of significant morphological differences" and considered "breedorum" invalid, noting "uncertain validity of this name resulting from publication practices." Gay refers to two femora of differing lengths (575, 605 mm) and two tibiae (560, 585 mm), believing more than one individual might be involved, but Welles states only the right hindlimb is preserved. Tykoski (2005) used the specimen as an example of D. wetherilli in his thesis and concluded it was an adult (unlike the types and TMM 43646) using an ontogenetic analysis. He states "at the behest of Kevin Padian (pers. comm., April, 2003) I refrain from giving a description of the crest morphology in the skull of UCMP 77270", perhaps indicating Padian or someone else is working on a new description of this specimen. Tykoski also states the quadratojugal, quadrate, sacral centra, distal pubis and metatarsals are not present in the specimen, which may mean they were lost after Welles' description. However, he notes a fibula is present, which is not mentioned by Welles. Tykoski further states the cervical ribs of UCMP 77270 are not fused to their vertebrae, contra Welles and Pickering. Irmis (2007) referred it to D. wetherilli and noted it had closed dorsal and proximal caudal neurocentral sutures, unlike the holotype. Carrano et al. (2012) noted the unfused interdental plates and trochanteric shelf differ from the holotype, but ascribed this to individual variation. Madsen and Welles (2000), Yates (2005), Sampson and Witmer (2007), Smith et al. (2007) and Carrano and Sampson (2008) all assigned it to D. wetherilli without comment.
Welles and Pickering diagnosed Dilophosaurus "breedorum" compared to D. wetherilli using several characters. Assessing their validity is made difficult by UCMP 77270 being older than the types (based on neurocentral fusion if nothing else) and some of Welles' (1984) description being based on casted features of the type patterned after Allosaurus. "Two, very thin, markedly developed parasagettal crests composed of the nasals + lacrimals + prefrontals" is also true in D. wetherilli, except that the participation of the prefrontals (on the medial surface) is uncertain due to UCMP 37302's crests being crushed together. Welles and Pickering later list the prefrontal participation as a separate autapomorphy. The dental formula only differs in having two more maxillary teeth and one less dentary tooth, which is usual individual variation in theropods. Welles and Pickering claim a separate postfrontal ossification is present in "breedorum", but state "the sutures are not obvious, and it could be absent or fused with the postorbital." Also they note the area in D. wetherilli is badly crushed, so this has little value as an apomorphy. The authors describe a deep groove along the posteroventral edge of the postorbital, which sounds similar to the condition in most megalosauroids (though I'm not sure if theirs also extends on to the posterior process). The quadratojugal has a posterolateral sulcus above the quadrate condyles and below the paraquadrate foramen. Both of these features are apparently different from the wetherilli holotype, but their significance is uncertain. Finally, Welles and Pickering state the cervical ribs are fused to their vertebrae (contra Tykoski), but this varies ontogenetically in "Megapnosaurus" kayentakatae anyway so would not be unexpected in an old Dilophosaurus individual. Being a Welles paper, most elements also include comparisons to their counterparts in D. wetherilli and other taxa. Most of these differences seem minor, though several support an adult stage of development- more elongate premaxilla, quadratojugal fused to quadrate, atlantal centrum and axial intercentrum fused to axis, deeper cervical pleurocoels, scapulocoracoid fusion, iliopubic fusion, trochanteric shelf present. Tykoski (2005) also noted other adult characters lacking in the holotype- various braincase and intersacral fusions, ilium fused to the sacrum, proximal femoral articular surfaces well developed, medial femoral epicondyle well developed, and an oblique ridge on the proximomedial fibula. Additionally, sacral central fusion was present as noted by Welles and Pickering. According to Tykoski, the interdental plates are unfused and tall in UCMP 77270 and TMM 43646, but that they are fused in UCMP 37302 and 37303. These differences cannot be explained by ontogeny and are not subject to individual variation in other taxa as far as I know. While they and some of the differences noted by Welles might suggest multiple species of Kayenta Dilophosaurus, other theropods known from large numbers of specimens (e.g. Allosaurus, Tyrannosaurus, Microraptor, Archaeopteryx) also show a high amount of morphological variation. I follow my recommendations for those taxa and only recognize a single species of Dilophosaurus, with UCMP 77270 simply being an older individual of D. wetherilli.
References- Welles, 1954. New Jurassic dinosaur from the Kayenta Formation of Arizona. Bulletin of the Geological Society of America. 65, 591-598.
Welles, 1970. Dilophosaurus (Reptilia: Saurischia), a new name for a dinosaur. Journal of Paleontology. 44, 989.
Welles, 1983. Two centers of ossification in a theropod astragalus. Journal of Paleontology. 57, 401.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica Abteilung A. 185, 85-180.
Clark and Fastovsky, 1986. Vertebrate biostratigraphy of the Glen Canyon Group in northern Arizona. The Beginning of the Age of the Dinosaurs: Faunal change across the Triassic-Jurassic boundary. in Fraser and Sues (eds.). Cambridge University Press. 285-301.
Gauthier, 1986. Saurischian Monophyly and the Origin of Birds. Memoires of the California Academy of Sciences. 8, 1-55.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Charig and Milner, 1990. The systematic position of Baryonyx walkeri, in the light of Gauthier's reclassification of the Theropoda. in Carpenter and Currie (eds.). Dinosaur Systematics: Approaches and Perspectives. Cambridge University Press, Cambridge. 127-140.
Rowe and Gauthier, 1990. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 151-168.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry. A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola, California. 478 pp.
Welles and Pickering, 1995. An extract from: Archosauromorpha: Cladistics and osteologies. A Fractal Scaling in Dinosaurology Project. 70 pp.
Olshevsky, DML online 1999. http://dml.cmnh.org/1999Dec/msg00097.html
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised osteology. Miscellaneous Publication 00-2, Utah Geological Survey. 80 pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.]. 440 pp.
Gay, 2001. Evidence for sexual dimorphism in the Early Jurassic theropod dinosaur, Dilophosaurus and a comparison with other related forms. Journal of Vertebrate Paleontology. 21(3), 53A.
Gay, 2001. New specimens of Dilophosaurus wetherilli (Dinosauria: Theropoda) from the Early Jurassic Kayenta Formation of northern Arizona. Western Association of Vertebrate Paleontologists annual meeting volume. Mesa, Arizona.
Gay, 2005. Sexual dimorphism in the Early Jurassic theropod Dilophosaurus and a comparison with other related forms. In Carpenter (ed.). The Carnivorous Dinosaurs. Indiana University Press. 277-283.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods. Palaeontologia Africana. 41, 105-122.
Mason, 2006. The thrill of the frill: Wear and tear demands restoration of a Dilophosaurus wetherilli skull, UCMP 77270. Journal of Vertebrate Paleontology. 26(3), 96A.
Irmis, 2007. Axial skeleton ontogeny in the Parasuchia (Archosauria: Pseudosuchia) and its implications for ontogenetic determination in archosaurs. Journal of Vertebrate Paleontology. 27(2), 350-361.
Sampson and Witmer, 2007. Craniofacial anatomy of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. in Sampson and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. SVP Memoir 8. 32-102.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Carrano and Sampson, 2008. The phylogeny of Ceratosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 6, 183-236.
Xu, Clark, Mo, Choiniere, Forster, Erickson, Hone, Sullivan, Eberth, Nesbitt, Zhao, Hernandez, Jia, Han and Guo, 2009. A Jurassic ceratosaur from China helps clarify avian digital homologies. Nature. 459, 940-944.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

unnamed clade (Liliensternus liliensterni + Coelophysis bauri)
= Coelophysidae sensu Sereno, 1998
Definition- (Coelophysis bauri + Procompsognathus triassicus) (modified)
Diagnosis- (after Tykoski, 2005) alveolar ridge on maxilla; deep ventral fossa on ectopterygoid; low post-axial cervical neural spines; ischiopubic ratio <66%; proximal end of metatarsal III with ventral boss; anterior ramus of maxilla <10% maxillary length; more than 19 maxillary teeth; anterior margin of antorbital fossa squared; anterior lacrimal process longer than ventral process; cervical ribs fused to vertebrae in adults.

"Comanchesaurus" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
"C. kuesi" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US
Material- (NMMNH P4569) dorsal centra, proximal femur, partial astragalus, metatarsal fragments, phalanges
Comments- NMMNH P4569 was originally seen as a coelophysoid by Lucas et al. (1985) and Hunt and Lucas (1989), though Murray and Long (1989) and Long and Murray (1995) referred it to Sauropodomorpha. Hunt (1994) named this "Comanchesaurus kuesi" in his unpublished thesis, assigning it to Herrerasauridae. It was called herrerasaurid B by Hunt et al. (1998). Nesbitt et al. (2007) confirmed a saurischian identity based on the dorsal astragalar basin, but noted the astragalar morphology was closer to coelophysids than to Herrerasaurus or other taxa. It may therefore be a coelophysoid, as originally believed.
Nesbitt et al. further noted the fragmentary remains referred to "Comanchesaurus" are indeterminate.
Although the name "Comanchesaurus kuesi" was originally used in thesis, and thus not available for use in this website, it was later published by Nesbitt et al. (2007).
References- Lucas, Hunt and Bennett, 1985. Triassic vertebrates from eastcentral New Mexico in the Yale Peabody Museum. New Mexico Geological Society Guidebook. 36, 199–203.
Hunt and Lucas, 1989. Late Triassic vertebrate localities in New Mexico. pp. 72-101. in Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque.
Murry and Long, 1989. Geology and paleontology of the Chinle Formation, Petrified Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils of the southwestern Upper Triassic. pp. 29-64. in Lucas & A. P. Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque.
Hunt, 1994. Vertebrate paleontology and biostratigraphy of the Bull Canyon Formation (Chinle Group: Norian), east-central New Mexico with revisions of the families Metoposauridae (Amphibia: Temnospondyli) and Parasuchidae (Reptilia: Archosauria). Unpublished PhD Dissertation. Albuquerque, Univerrsity of New Mexico. 403 pp.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

Dolichosuchus Huene, 1932
D. cristatus Huene, 1932
Early-Middle Norian, Late Triassic
Lower or Middle Stubsandstein, Germany
Holotype- (BMNH R38058) tibia (330 mm)
Diagnosis- Provisionally indeterminate relative to Liliensternus liliensterni.
Comments- Huene (1932) originally assigned this genus to Hallopodidae.
This has a large cnemial crest and fibular crest, showing it is theropod. Welles (1984) found the differences from Liliensternus to be insignificant. Rauhut and Hungerbuhler (2000) note close resemblence to Liliensternus and Dilophosaurus, suggesting it is probably a coelophysoid.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monog. Geol. Pal. 4 (1) pts. 1 and 2, viii + 361 pp.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie 185 p. 85-180.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.

Procompsognathidae Nopcsa, 1923 sensu Huene, 1929
Comments- Huene (1929) separated Procompsognathus from podokesaurids as a new family Procompsognathidae, which was not followed by many authors through the 1900's. Huene (1932) included Pterospondylus as well, which was often viewed as a synonym of Procompsognathus. Romer (1966) and others have incorrectly used this as a senior synonym of Podokesauridae, including Avipes, Coelophysis, Dolichosuchus, Halticosaurus, Lukousaurus, Podokesaurus, Procompsognathus, Saltopus, Scleromochlus, Spinosuchus, Trialestes and Velocipes in the family. Steel (1970) only included Procompsognathus and Halticosaurus. Welles (1984) and Paul (1988) both only included Procompsognathus in the family. Procompsognathidae has not been used since the 1980's, as Procompsognathus has most often been viewed as a coelophysoid incertae sedis or coelophysid, and was even made an internal specifier of Coelophysidae by Sereno (1998). Yet if the genus is a basal dinosauriform as Allen (2004) believes, the family may be justified again.
References- Nopcsa, 1923. Die Familien der Reptilien [The families of reptiles]. Forschritte der Geologie und Palaeontologie. 2, 1-210.
Huene, 1929. Kurze Übersicht über die Saurischia und ihre natürlichen Zusammenhänge [A brief survey of the Saurischia and their natural context]. Paläontologische Zeitschrift. 11, 269-273.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monog. Geol. Pal. 1, pts 1 and 2, V111 + 361 pp.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press, Chicago. 1-468.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 1-87.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 210(1), 41-83.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP 24(3).
Procompsognathinae Nopcsa, 1923
Definition- (Procompsognathus triassicus <- Coelophysis bauri) (Sereno, in press; modified from Sereno, 1998)
Comments- Nopcsa (1923) named this subfamily within Compsognathidae, separate from Compsognathinae and Ornithomiminae. This was not followed, though Sereno (1998) later used it as a subfamily of Coelophysidae to contain Procompsognathus and Segisaurus, separate from the coelophysines Coelophysis and Megapnosaurus. This was not found in future studies (e.g. Tykoski, 2005; Ezcurra and Novas, 2006), where Procompsognathus instead has a more unstable position more derived than Dilophosaurus. This makes the utility of Procompsognathinae questionable and the application of Sereno's definition to any taxon except Procompsognathus itself impossible. If Procompsognathus is non-dinosaurian as Allen (2004) suggests, Procompsognathinae may have some use depending on its exact relationships.
References- Nopcsa, 1923. Die Familien der Reptilien [The families of reptiles]. Forschritte der Geologie und Palaeontologie. 2, 1-210.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 210(1), 41-83.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP 24(3).
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article, 38 pp. DOI 10.1080/08912960600845791.
Procompsognathus Fraas, 1913
P. triassicus Fraas, 1913
= Hallopus "celerrimus" Fraas, 1912
Middle Norian, Late Triassic
Middle Stubensandstein, Germany
Holotype- (SMNS 12591) (.96 m) (robust) skull (associated?), partial cervical vertebrae 5-10, partial cervical ribs, first dorsal vertebra, second dorsal vertebra (~12.6 mm), third dorsal vertebra (~13.6 mm), fourth dorsal vertebra (~13.6 mm), fifth dorsal vertebra, sixth dorsal vertebra (15.8 mm), seventh dorsal vertebra (16.2 mm), eighth dorsal vertebra (16.2 mm), ninth dorsal vertebra (16.8 mm), tenth dorsal vertebra (16.4 mm), partial dorsal ribs, thirteen anterior caudal vertebrae 15 mm), scapulocoracoid (43+ mm), radius, ulna (34.2 mm), radiale?, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, partial phalanx II-1, incomplete metacarpal III, partial metacarpal IV, partial ilium, pubes (88+, 86+ mm), femora (92.5, 93.1 mm), tibia (112.6 mm), proximal tibia, fibula, astragalus, metatarsal I (10.7 mm), phalanx I-1 (12.3 mm), pedal ungual I (7.3 mm), metatarsal II (58 mm), phalanx II-1 (15.9 mm), phalanx II-2 (12.8 mm), pedal ungual II (10.6 mm), metatarsal III (69.4 mm), phalanx III-1 (17.1 mm), phalanx III-2 (15.2 mm), phalanx III-3 (12 mm), pedal ungual III (9.7 mm), metatarsal IV (68.8 mm), phalanx IV-1 (~6.8 mm), phalanx IV-2 (8.7 mm), phalanx IV-3 (7.2 mm), phalanx IV-4 (5.2 mm), pedal ungual IV (7.2 mm), metatarsal V (23 mm) (Huene 1921, Ostrom 1981, Sereno and Wild 1992, Chatterjee 1993)
Diagnosis- (after Rauhut, 2000) scapula more slender than Coelophysis bauri.
Comments- Hallopus "celerrimus" is probably a synonym of Procompsognathus triassicus, according to Molnar (pers. comm.). He suspects this based on the fact they come from the same locality and beds, were proposed by the same person, and Fraas never used the name Hallopus "celerrimus" after he published Procompsognathus triassicus.
Sereno and Wild (1992) considered the skull to belong to Saltoposuchus connectens, while Chatterjee (1993) believed it to be theropod though the latter evidence is not published yet.
A detailed description of the reprepared skull is in progress by Chatterjee (pers. comm. to Rauhut, 1996), so Rauhut and Hungerbuhler (2000) do not consider the skull in their commentary. They note that the slightly bowed pubis with rectangular apron and absence of a pubic foot in this specimen are symplesiomorphies, so do not help resolve relationships (contra Ostrom 1981, Sereno and Wild 1992). They think the elongate dorsal vertebrae and triangular dorsal transverse processes may indicate ceratosaurian or coelophysoid affinities. It is provisionally regarded as valid based on an elongate hindlimb (tib/fem 1.2, mtIII/fem .74). I do not find this character convincing, as Podokesaurus has similar ratios (tib/fem 1.21, mtIII/fem .76) and Coelophysis can also have similar proportions (tib/fem 1.17, mtIII/fem .71). Still, I feel there is a good possibility future work will provide valid synapomorphies.
Allen (2004) believes the skull belongs to the same taxon as the postcrania, and based on cladistic analyses both including and excluding the former, places Procompsognathus as a non-dinosaurian avemetatarsalian. However, the evidence for this reinterpretation has not yet been published.
References- Fraas, 1912. Jahresh. Ver. Naturk. Wuerttemberg. 68:66-67.
Fraas, 1913. Die neuesten Dinosaurierfunde in der schwabischen Trias. Naturwissenschaften 45: 1097-1100.
Ostrom, 1981. Procompsognathus - theropod or thecodont?. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie 175 p. 179-195.
Sereno and Wild, 1992. Procompsognathus: Theropod, ``thecodont'' or both?. Journal of Vertebrate Paleontology 12 p. 435-458.
Chatterjee, 1993. Procompsognathus from the Triassic of Germany is not a crocodylomorph. Abstract, Journal of Vertebrate Paleontology, 13(3): 29A.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP 24(3).

Protoaviformes Chatterjee, 1991
= Praeornithurae Kurochkin, 1995
= Protoaviornithes Kurochkin, 1995
Protoavidae Chatterjee, 1991
Protoavis Chatterjee, 1991
P. texensis Chatterjee, 1991
Norian, Late Triassic
Bull Canyon Formation of Dockum Group, Texas, US
Holotype- (TTU P 9200)
unassigned- premaxilla, lacrimal(?), squamosal, quadrate(?), frontal, parietal, posterior mandible(?), atlas(?), axis (10 mm), two caudal vertebrae (10 mm), partial clavicle(?) (49 mm), distal scapula(?) (~37 mm), ilia(?) (43 mm), partial pubis(?) (37 mm), ischia(?) (22 mm)
Drepanosauridae- cervical vertebra (18 mm)
Pterosauromorpha- ischium
Theropoda- basioccipital, prootic, exoccipital, epiotic, supraoccipital
Coelophysoidea- (juvenile?) incomplete femur (~58 mm)
Paratype- (TTU P 9201)
unassigned- maxilla, nasal(?), lacrimal, jugal, quadratojugal, quadrate(?), parietal(?), basioccipital(?), vomer, palatine, pterygoid, partial dentary, posterior dentary(?), posterior mandible, axis (8 mm), cervical rib, seven dorsal vertebrae (7-10 mm), two dorsal ribs, two sacral vertebrae (7 mm), eighteen caudal vertebrae (7-9 mm), two chevrons, scapula(?) (~21 mm), coracoid(?) (18 mm), humerus (~39 mm), ilium(?) (22 mm), proximal tibia, proximal fibulae, metatarsal I(?) (9 mm), phalanx (10 mm), phalanges, unguals
Drepanosauridae- seven cervical vertebrae (9-12 mm)
Pterosauromorpha- tibia (~39 mm), fibula (~41 mm), four tarsals, metatarsal I (12 mm), metatarsal II (20 mm), metatarsal III (18 mm), metatarsal IV (14 mm)
Theropoda- distal tarsal IV, metatarsal II (25 mm), metatarsal III (27 mm), metatarsal IV (25 mm)
Coelophysoidea- (juvenile?) astragalus, calcaneum
Late Carnian, Late Triassic
Tecovas Formation of the Dockum Group, Texas, US
Referred- (TTU P 9350-9355) six dorsal vertebrae (6-10 mm) (Chatterjee, 1995)
(TTU P 9356-9359) four caudal vertebrae (8-9 mm) (Chatterjee, 1995)
(TTU P 9360) coracoid(?) (14 mm) (Chatterjee, 1995)
(TTU P 9361) sternum(?) (25 mm) (Chatterjee, 1995)
(TTU P 9362) humerus (~80 mm) (Chatterjee, 1995)
(TTU P 9263) humerus (Chatterjee, 1995)
(TTU P 9364) partial mandible (Chatterjee, 1995)
(TTU P 9365) humerus (Chatterjee, 1995)
(TTU P 9367) pterosauromorph tibia (Chatterjee, 1995)
(TTU P 9368) pterosauromorph tibia (~58 mm) (Chatterjee, 1995)
(TTU P 9369) pterosauromorph fibula (Chatterjee, 1995)
(TTU P 9370) femur (~58 mm) (Chatterjee, 1995)
(TTU P 9371) femur (Chatterjee, 1995)
(TTU P 9372) femur (Chatterjee, 1995)
(TTU P 9373) femur (Chatterjee, 1995)
(TTU P 9374) tibia (Chatterjee, 1995)
(TTU P 9375-9380) phalanges (Chatterjee, 1995)
Diagnosis- A chimaerical taxon whose holotype includes portions of multiple organisms.
Comments- Initially identified as a juvenile Coelophysis (Chatterjee, 1986), this material was then reported as the oldest bird (Wilford, 1986; Chatterjee, 1987, 1988). This was met with skepticism from the paleontological community, with Ostrom (1987) questioning its avialan identification, while Paul (1988) tentatively believed it to be a herrerasaurian.
After its official description was published (Chatterjee, 1991), some agreed it was avialan (Kurochkin, 1992; Peters, 1994; Kurochkin, 1995; Martin, 2004). When analyzed as a bird, Protoavis is resolved as- an ornithurine intermediate between Avimimus and Ornithothoraces (Chatterjee, 1991; Chatterjee, 1998; Dyke and Thorley, 1998); or an enantiornithine-grade ornithothoracine (Chatterjee, 1999). Kurochkin (1995) hypothesized it was the basalmost ornithurine (sensu Feduccia, Martin, etc.).
However, most dismissed an avialan identity (Ostrom, 1991; Wellnhofer, 1992; Chiappe, 1995; Feduccia, 1996; Ostrom, 1996; Sereno, 1997; Hunt et al., 1998; Renesto, 2000; Nesbitt et al., 2005). These authors have usually suggested the remains are chimaerical, including a non-avian coelurosaur braincase (Witmer, 2001), megalancosaurid cervicals (Renesto, 2000), possibly lepidosauromorph humerus (Witmer, 2001), coelophysoid femur and proximal tarsals (Hutchinson, 2001; Nesbitt et al., 2005), and non-avian archosaurian pes (Sereno, 1997). Chinle and Dockum megalancosaurids (Dolabrosaurus; Harris and Downs, 2002), lepidosauromorphs (Clevosaurus), coelophysoids (Coelophysis, Gojirasaurus) and crurotarsans (parasuchians, aetosaurs, poposaurids, rauisuchids) are known, but if some elements are coelurosaurian, it would be unprecedented.
Witmer (2001) has undertaken the most detailed independant analysis of the remains, and concludes many of the morphologies identified by Chatterjee (e.g. pterygoid cotyla on quadrate; mandibular condyles on quadrate; quadrate foramina; quadratojugal cotyle on quadrate; basisphenoid; scapulae; clavicle; coracoidal sulci on sternum; intermuscular line on sternum; ulnar quill knobs; metacarpal quill knobs; opisthopuby; fused ilium and ischium; ischial antitrochanter; tibiotarsal fusion) cannot be confirmed. However, he notes several characters are birdlike- apparently absent contact between squamosal and quadratojugal and postorbital; heterocoelous cervical vertebrae; well developed cervical hypapophyses; large cervical neural canals; coracoid morphology (elongated; strut-like; procoracoid and acrocoracoid processes).
Witmer described the large floccular fossa, cranial pneumatic recesses and metotic strut as coelurosaurian characters in the holotype braincase. However, the metotic strut is a tetanurine character, the anterior and dorsal tympanic recesses are theropod characters, the posterior tympanic recess has been described in Ceratosaurus, and large floccular fossae are known at least as basally as Ceratosauria. Furthermore, Protoavis' floccular fossa appears to be narrow like non-maniraptorans, and its posterior temporal recess opens outside the otic recess unlike avialans. So while it's possible the braincase belongs to a tetanurine or ceratosaur, more information on coelophysoid braincase variation and ontogeny would be ideal.
Drepanosaurids share the avian cervical characters noted above with Protoavis and the vertebrae of the two taxa are extremely similar, making it highly likely some material is referrable to Dolabrosaurus or another drepanosaurid.
Hutchinson (2001) noted the proximal femur is identical to coelophysid femora and suggested it may be a juvenile. Witmer (2001) found the proximal tarsals to resemble non-coelurosaurian theropods, while Nesbitt et al. (2005) confirmed all these elements are coelophysoid. Nesbitt et al. (2007) noted the following dinosaur and theropod characters in the femur- offset head; ligament sulcus; trochanteric fossa; trochanteric shelf; small posterior trochanter. They agreed with Hutchinson it's similar to coelophysoids, but found no apomorphies of that clade, and noted the fourth trochanter is not absent (contra Chatterjee, 1999), but that the area is unpreserved. Nesbitt et al. also noted coelophysoid-like tarsal characters, such a a fibular facet formed from both proximal tarsals (unlike neoceratosaurs and tetanurines) and the distally (not anteriorly) articulated proximal tarsals (unlike tetanurines). It's probable these belong to juvenile Coelophysis (as originally believed) or Gojirasaurus. Additional Coelophysis-like material is known from the Dockum Group (UCMP online; Lehman and Chatterjee, 2005). The astragalus' low ascending process indicates it is a coelophysoid more closely related to Coelophysis than Dilophosaurus.
Many elements of Protoavis appear to be from a new supposed pterosauromorph named and described in a thesis (Atanassov, 2001, 2002). The original (Chatterjee, 1991; reidentified as a vomer and pterygoid in 1999) sternum is an ischium, the radius and ulna are the tibia and fibula, and the metacarpus is the metatarsus. The new form's maxilla, dentary, ilium and femur seem unrepresented in Protoavis' material, while some characters of the vertebrae are a good match. Unfortunately, only two vertebrae (both sacrals) were associated with the appendicular and cranial elements of this new taxon, so it's possible the latter's vertebrae are from a drepanosaurid or other taxon.
References- Chatterjee, 1986. The Late Triassic Dockum vertebrates: Their stratographic and paleobiogeographic significance. In Padian (ed.). The beginning of the age of dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge University Press. 378 pp.
Wilford, 1986. Texas fossil may be bird's oldest ancestor. New York Times. 8-14-1986.
Chatterjee, 1987. Skull of Protoavis and early evolution of birds. JVP. 7(3) 14A.
Ostrom, 1987. Protoavis, a Triassic bird?. Archaeopteryx. 5, 113-114.
Chatterjee, 1988. Functional significance of the semilunate carpal in archosaurs and birds. JVP. 8(3) 11A.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Ostrom, 1991. The bird in the bush. Nature. 353, 212.
Chatterjee, 1991. Cranial Anatomy and relationships of a new Triassic bird from Texas. Philosophical Transactions of the Royal Society of London Series B 332(1265): 277-342.
Kurochkin, 1991. Protoavis, Ambiortus and other paleornithological rarities. [in Russian]. Priroda. 1991, 43-53.
Chatterjee, 1992. Texas fossil identified as the world's oldest known bird. J. Geol. Soc. India 39 p. 89-90.
Viohl, 1992. No evidence of Triassic birds. [in German]. Archaeopteryx. 10, 77-79.
Wellnhofer, 1992. Protoavis: The oldest bird?. [in German]. Naturwiss. Rundsch. 45, 107-108.
Kurochkin, 1992. The oldest bird. [in Russian]. Priroda. 1992, 100-101.
Peters, 1994. Die Entstehung der Vogel-Verandern die jungsten Fossilfunde das Modell? In Gutmann et al. (eds.). Morphologie und evolution. Frankfurt. 403-424.
Chatterjee, 1995. The Triassic bird Protoavis. Archaeopteryx. 13, 15-31.
Chiappe, 1995. The first 85 million years of avian evolution. Nature. 378, 349-355.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of Class Aves. Archaeopteryx. 13, 47-66.
Feduccia, 1996. The Origin and Evolution of Birds. Yale University Press. 420 pp.
Ostrom, 1996. The Questionable Validity of Protoavis. Archaeopteryx. 14, 39-42.
Chatterjee, 1997. The Rise of Birds: 225 Million Years of Evolution. John Hopkins University Press. 312 pp.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences. 25, 435-489.
Chatterjee, 1998. The avian status of Protoavis. Archaeopteryx. 16, 99-122.
Dyke and Thorley, 1998. Reduced cladistic consensus methods and the inter-relationships of Protoavis, Avimimus, and Mesozoic birds. Archaeopteryx. 16, 123-129.
Hunt, A.P., S.G. Lucas, A.B. Heckert, R.M. Sullivan and M.G. Lockley. 1998. Late Triassic dinosaurs from the western United States. Geobios, 31(4):511-531.
Chatterjee, 1999. Protoavis and the early evolution of birds. Palaeontographica A. 254, 1-100.
Renesto, 2000. Bird-like head on a chameleon body: new specimens ofthe enigmatic diapsid reptile Megalancosaurus from the Late Triassic of northern Italy. Rivista Italiana di Paleontologia e Stratigrafia. 106:157–180.
Atanassov, 2001. Two new archosauromorphs from the Late Triassic of Texas. Journal of Vertebrate Paleontology. 21(3) 30A.
Hutchinson, 2001. The evolution of femoral osteology and soft tissues on the line to extant birds (Neornithes). Zoological Journal of the Linnaean Society. 131, 169-197.
Witmer, 2001. The role of Protoavis in the debate on avian origins. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds. Yale University. 538-548.
Atanassov, 2002. Unpublished thesis. Texas Tech University.
Harris and Downs, 2002. A drepanosaurid pectoral girdle from the Ghost Ranch (Whitaker) Coelophysis Quarry (Chinle Group, Rock Point Formation, Rhaetian), New Mexico. JVP. 22(1), 70-75.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica. 50(6), 978-990.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. JVP 25(3) 96A.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

Liliensternus Welles, 1984
Diagnosis- (after Rauhut, 2000) broad rounded ridge on the cervical vertebrae, that extends from the posterior end of the diapophyses to the posteroventral end of the vertebral centrum.
L. liliensterni (Huene, 1934) Welles, 1984
= Halticosaurus liliensterni Huene, 1934
Late Norian, Late Triassic
Trossingen Formation, Germany
Syntypes- (MB.R.2175; = HMN R1291) (gracile young subadult) (5.15 m, 127 kg) (partial skull 395 mm), ventral maxilla, jugal, postorbital, incomplete squamosal, quadrate, dentary, surangular, partial angular, fourth cervical vertebra (70 mm), tenth cervical vertebra (47 mm), first sacral vertebra (52 mm), second sacral vertebra (52 mm), third sacral vertebra (52 mm), scapula (~240 mm), humerus (214 mm), radius (151 mm), ulna (158 mm), metacarpal II (70 mm), ilium (270 mm), pubis (418 mm), ischium (303 mm), femur (440 mm), tibia (409 mm), astragalocalcaneum (79 mm wide, astragalus 60 mm wide), metatarsal II (214 mm), phalanx II-1 (73 mm), phalanx II-2 (39 mm), pedal ungual II (55 mm), metatarsal III (230 mm), phalanx III-1 (69 mm), phalanx III-2 (55 mm), phalanx III-3 (46 mm), pedal ungual III (46 mm), metatarsal IV (221 mm), phalanx IV-1 (51 mm), phalanx IV-2 (35 mm), phalanx IV-3 (29 mm), phalanx IV-4 (27 mm)
(MB.R.2175) (gracile young subadult) partial skeleton
Diagnosis- (after Rauhut, 2000) one pair of pleurocoels in the cervical vertebrae, less developed infradiapophyseal fossa; the absence of a horizontal ridge at the basis of the cervical neural spines; absence of a lateral bulge on the ilium.
Comments- The syntype remains of this species are usually referred to two individuals. However, the material was found disarticulated and may represent more than two individuals. Because of this and the fact it is hard to separate the remains belonging to the various individuals, Rauhut and Hungerbuhler (2000) recommend retaining all of the material as the syntypes of Liliensternus (contra Welles 1984, who made the larger individual the syntype). Because the neurocentral sutures are unfused and only two fused sacrals are present, the remains are probably juveniles or subadults. Contra Rowe and Gauthier (1990), the pubis encloses a complete obturator foramen, not just a notch.
References- Huene, 1934. Ein neuer Coelurosaurier in der thuringischen Trias. Pal. Zeit. 16 145-170, 4 pls.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie 185 p. 85-180.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.
L? sp. indet. (Sander, 1992)
Late Norian, Late Triassic
Trossingen Formation, Germany
Material- (UA coll.) proximal metatarsal II (Galton, 2000)
at least two occurances (Sander, 1992)
Comments- The UA coll. metatarsal was originally a syntype of Plateosaurus engelhardti, described as a manual or pedal element by Meyer (1855) and a pubic fragment by Huene (1908). It was reidentified as a proximal metatarsal IV by Galton (2000) and a proximal metatarsal II by Moser (2003). Comparison with figures of Liliensternus indicates Moser is correct.
Reference- Meyer, 1955. Zur fauna der Vorwelt. Die saurier des Muschelkalkes, rnit Rucksicht auf die saurier aus Buntem Sandstein und Keuper. 167 pp.
Huene, 1908. Die Dinosaurier der Europaiaschen Triasformation mit Berucksichtiging der aussereuropaischen Vorkommnisse. Geol. Paleont. Abhandl. Suppl. 1, 1-419.
Sander, 1992. The Norian Plateosaurus bonebeds of central Europe and their taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology. 93:255-296.
Galton, 2000. The prosauropod dinosaur Plateosaurus Meyer, 1837 (Saurischia: Sauropodomorpha). I. The syntypes of P. engelhardti Meyer, 1837 (Upper Triassic, Germany), with notes on other European prosauropods with "distally straight" femora. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen. 216(2): 233-275.
Moser, 2003. Plateosaurus engelhardti Meyer, 1837 (Dinosauria: Sauropodomorpha) aus dem Feuerletten (Mittelkeuper; Obertrias) von Bayern. Zitteliana B 24, 3-186.
L? sp. indet. (Sander, 1992)
Norian, Late Triassic
Knollenmergel, Switzerland
Reference- Sander, 1992. The Norian Plateosaurus bonebeds of central Europe and their taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology. 93:255-296.

unnamed clade (Zupaysaurus rougieri + Lophostropheus airelensis + Coelophysis bauri)
Diagnosis- (modified from Tykoski, 2005) astragalus and tibia fused in adults.
(after Ezcurra and Novas, 2006)- angle between the rostrodorsal margin of the maxilla and the alveolar margin less than 75 degrees (unknown in Lophostropheus); infratemporal fenestra strongly rostrocaudally compressed, with maximum length versus maximum length of the orbit ratio less than 0.8 (unknown in Lophostropheus); caudal curvature of the dorsal end of quadrate, with quadrate head caudodorsally oriented (unknown in Lophostropheus).
(after Ezcurra and Cuny, 2007) supraacetabular crest continuous with the lateral margin of the brevis fossa, forming a well-developed ridge, without a notch between both structures (unknown in Zupaysaurus).

Lophostropheus Ezcurra and Cuny, 2007
L. airelensis (Cuny and Galton, 1993) Ezcurra and Cuny, 2007
= Liliensternus airelensis Cuny and Galton, 1993
Late Rhaetian-Early Hettangian, Late Triassic-Early Jurassic
Moon-Airel Formation, France
Holotype- (Caen University coll.) tooth, five cervical vertebrae (72, 83 mm), two posterior dorsal vertebrae (75 mm), four sacral vertebrae, several caudal vertebrae (72 mm), partial ilium, proximal pubes, partial ischium
Diagnosis- (after Rauhut, 2000) deep infradiapophyseal fossa in anterior cervical vertebrae; horizontal ridge at the basis of the neural spine in cervical vertebrae; ilium with a triangular lateral bulge above the supraacetabular crest.
(after Ezcurra and Cuny, 2007) moderately convex anterior articular surface of the anterior postaxial cervical vertebrae (also in Ceratosauria+Tetanurae); large and
oval lateral fossa on last dorsal vertebral centrum (also in Herrerasaurus); dorsoventrally well-extended hyposphene in the last dorsal vertebra; incipient concavity on the cranial articular surface of cranial caudal vertebrae (also present in Ceratosauria+Tetanurae); constant length of caudal vertebrae along the tail (also in Dilophosaurus).
Comments- Rauhut and Hungerbuhler (2000) also listed the "cervical vertebrae with dorso-ventrally narrow, antero-posteriorly elongated posterior pleurocoel" as being diagnostic, but it is also present in Coelophysis.
Although Rauhut and Hungerbuhler (2000) believe this is a valid species, they think characters such as the presence of two pairs of cervical pleurocoels (versus one pair in L. liliensterni), may indicate it deserves a separate genus.
References- Larsonneur and Lapparent, 1966. Un dinosaurien carnivore, Halticosaurus, dans le Réthien d´ Airel (Manche). Bulletin Societe Linneenne de Normandie. 10:108–116.
Cuny and Galton, 1993. Revision of the Airel theropod dinosaur from the Triassic-Jurassic boundary (Normandy, France). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 187 p. 261-288.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.
Ezcurra and Cuny, 2007. The coelophysoid Lophostropheus airelensis, gen. nov.: Areview of the systematics of "Liliensternus" airelensis from the Triassic-Jurassic boundary outcrops of Normandy (France). Journal of Vertebrate Paleontology. 27(1), 73-86.

Zupaysaurus Arcucci and Coria, 2003
= “Zupaysaurus” unpublished, 1999
Z. rougieri Arcucci and Coria, 2003
Rhaetian, Late Triassic
Upper Los Colorados Formation, La Rioja, Argentina
Holotype- (PULR-076) (5-6 m) skull (450 mm), mandibles, atlas, axis (110 mm), cervical vertebrae, cervical ribs, dorsal vertebrae, sacral vertebrae, distal caudal vertebrae, proximal scapulocoracoid, two proximal manual unguals, distal femora, proximal tibia, distal tibia, distal fibula, astragalocalcaneum
Diagnosis- (modified from Arcucci and Coria, 2003) horizontal ramus of the maxilla with parallel dorsal and ventral margins; tibia with a very deep and caudally open notch for the reception of an astragalar caudal process.
(after Ezcurra, 2006) maxillary fenestra within the antorbital fossa (also in Tetanurae); ventrally bowed rostral process of the lacrimal (also in Sinraptor); kinked ventral process of the squamosal; wide contact between squamosal and quadratojugal (also in several tetanurines).
(after Ezcurra and Novas, 2006) maxillary-jugal ventral margin describing an obtuse angle in lateral view; notch on the dorsal margin of the ascending process of the maxilla, relating to horizontal ramus of lacrimal rostrally tapering onto the forked caudal tip of the ascending process of the maxilla; lacrimal with highly pneumatized antorbital recess (also in Ceratosauria+Tetanurae); short and square-shaped retroarticular process of the mandible; cnemial crest poorly developed.
Comments- Originally thought to be "more derived than Coelophysis" (Arcucci and Coria, 1997), and later a ceratosaur (Arcucci and Coria, 1998). When officially described by Arcucci and Coria (2003), it was found to be a basal tetanurine. However, later analyses have found that it is in fact a coelophysoid. Carrano et al. (2005) found it to be a coelophysoid, but could not recover any resolution within that clade. Zupaysaurus was found to be in a trichotomy with Liliensternus and coelophysids by Tykoski (2005). Ezcurra and Novas (2005) agree the taxon is a coelophysoid, which was elaborated on in Ezcurra and Novas (2006). Of the supposed tetanurine characters, some are reinterpreted as convergences (maxillary fenestra; lacrimal recess; fibula with distal end expanded almost double the shaft width), while others aren't present in Zupaysaurus (antorbital maxillary tooth row; lacrimal horn; distal tibia transversely expanded) or are present in coelophysoids as well (lateral temporal fenestra reduced and key-hole-shaped; tibia with a caudolaterally concave distal end; ascending process of astragalus cranially positioned). They found it to be in a trichotomy with Segisaurus and Coelophysidae. Smith et al. (2007) found Zupaysaurus to be more derived than coelophysoids, sister to a clade containing dilophosaurids, ceratosaurs and tetanurines. This was based on- tooth row that ends at the anterior rim of the orbit (not actually present); jugal with an expanded anterior end; lacrimal fenestra; broad contact between the squamosal and quadratojugal; well-developed anterior wall to the lateral mandibular glenoid; broadened retroarticular process; posterodorsally facing surface for the attachment of the m. depressor mandibulae on the retroarticular process; astragalar ascending process that is higher than the astragalar body. Placing Zupaysaurus inside Coelophysoidea took five more steps.
References- Arcucci and Coria, 1997. First record of Theropoda (Dinosauria - Saurischia) from the Los Colorados Formation (Upper Triassic, La Rioja, Argentina). Ameghiniana 34(4) 531.
Arcucci and Coria, 1998. Skull features of a new primitive theropod from Argentina. JVP 18(3) 24A-25A.
unpublished, 1999. Discovery News. 11-12-1999.
Arcucci and Coria, 2003. A new Triassic carnivorous dinosaur from Argentina. Ameghiniana 40(2): 217 - 228.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate Paleontology. 25(4), 835–849.
Ezcurra and Novas, 2005. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. In Kellner, Henriques and Rodrigues (eds.), II Congresso Latino-Americano de Paleontologia de Vertebrados, Boletim de Resumos. Museum Nacional/UFRJ, Rio de Janeiro 102-104.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. The cranial anatomy of the coelophysoid theropod Zupaysaurus rougieri from the Upper Triassic of Argentina. Historical Biology. iFirst Article, 18 pp. DOI 10.1080/08912960600861467.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article, 38 pp. DOI 10.1080/08912960600845791.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.

Coelophysidae Nopcsa, 1928 sensu Welles, 1984
Definition- (Coelophysis bauri + Megapnosaurus rhodesiensis + "Megapnosaurus" kayentakatae) (Holtz, 1994)
Other definitions- (Coelophysis bauri + Procompsognathus triassicus) (Sereno, in press; modified from Sereno, 1998)
(Coelophysis bauri + Megapnosaurus rhodesiensis) (modified from Tykoski and Rowe, 2004)
= Podokesauridae Huene, 1914a
= Podokesaurinae Huene, 1914a sensu Nopcsa, 1923
= Segisauridae Camp, 1936
Diagnosis- (modified from Tykoski, 2005) medial flange of fibula overlaps astragalar ascending process; fibular facet of astragalus reduced, subtriangular in shape on anterolateral corner of proximal surface; postaxial cervical neural arches house pneumatic cavities lateral to neural canal; lateral surface of postaxial cervical neural arch pediucels have posteriorly directed fossae anterior to postzygapophyses; elongate mid cervical centra (>3 times height of anterior articular surface); elongate posterior dorsal centra (>2 times height of anterior articular surface); m. iliofemoralis fossa of ilium stops short of posterior margin, leaving distinct lateral rim on postacetabular process; distal tarsal III fuses to metatarsal III in adults; metatarsals II and III fuse in adults; circular orbit; few or no neurovascular foramina on lateral premaxillary surface; medial surface of anteromedial maxillary process smooth; low rounded horizontal ridge on lateral surface of jugal; ventral process of squamosal narrow; lamina spanning the basal tubera bears small median spur that projects anteriorly along roof of basisphenoid recess; anterior premaxillary teeth subcircular in section; anterior premaxillary teeth with reduced curvature; cervical ribs elongate (>4 times central length); posterior margin of ilium concave in lateral view.
Comments- Tykoski (2005) also included the presence of a furcula as an ambiguous synapomorphy of this clade, possibly based on a miscoding of Carnotaurus.
Podokesauridae- Podokesauridae was first proposed by Huene (1914a) as a basal grade of coelurosaurs (sensu Huene). This position was retained throughout the mid-1900's, with podokesaurids seen as directly ancestral to coelurids, and less directly to compsognathids and ornithomimids. Barsbold (1977) had podokesaurids ancestral to all theropods, and coelurids and troodontids in particular. Huene (1914) originally included Podokesaurus, Coelophysis, Saltopus and Tanystropheus. Later added were Procompsognathus (Huene, 1914b), Halticosaurus (Huene, 1914b), Procerosaurus (Huene, 1920), Avipes (Huene, 1932), Spinosuchus (Huene, 1932), Dolichosuchus (Romer, 1956), Lukousaurus (Romer, 1956), Velocipes (Romer, 1956), Trialestes (Reig, 1963; as Triassolestes), Megapnosaurus (Raath, 1969; as Syntarsus), Dilophosaurus (Russell, 1984), Alwalkeria (Chatterjee, 1987; as Walkeria), Segisaurus (Carroll, 1988), "Comanchesaurus" (Hunt and Lucas, 1989), Sarcosaurus (Madsen and Welles, 2000) and Liliensternus (earlier as Halticosaurus liliensterni, then explicitly by Madsen and Welles, 2000). The family was generally not used after the 1980's, with its genera then referred to as ceratosaurs, coelophysoids and/or coelophysids. Madsen and Welles (2000) correctly noted that Podokesauridae has priority over Coelophysidae according to the ICZN, since it was named fourteen years earlier. Yet all other workers ignore this, generally because they value Phylocode rules and Podokesauridae has not been phylogenetically defined, or they misread the ICZN to demand family-level eponyms be diagnosable (it actually only says they must be valid). Coelophysidae is only used on this website because of its near universal usage since 1991.
Nopcsa (1923) changed it to subfamily Podokesaurinae within Hallopodidae, which has not been followed as Hallopus is a crocodylomorph.
Coelophysidae- Welles (1984) erected Coelophysidae from Nopcsa's earlier subfamily Coelophysinae, including Coelophysis and Sarcosaurus without comment. Paul (1988) later included Coelophysis, Dilophosaurus, Elaphrosaurus, Liliensternus and Megapnosaurus (as Coelophysis rhodesiensis) in the family. This was followed by Novas (1991, 1992), though Holtz (1994) restricted the term to exclude the basal Dilophosaurus. This is followed by all workers currently yet the precise extent of Coelophysidae has since been controversial, sometimes including Liliensternus (Carrano et al., 2002), and sometimes restricted to Coelophysis and Megapnosaurus (Rauhut, 2003). This depends both on the definition used and the resolution within derived Coelophysoidea. An intermediate extent is used here, based on the definition in Holtz (1994) and the analysis of Tykoski (2005).
Segisauridae- Segisauridae was established by Camp (1936) to include only Segisaurus, viewed as related to coelurids and compsognathids within Coelurosauria sensu Huene. This practice was followed until Gauthier (1986) determined that Segisaurus was a ceratosaur sensu lato. Segisauridae has been almost unused since, with Segisaurus being placed in Coelophysoidea and/or Coelophysidae instead (Sereno, 1997). If Coelophysidae is restricted to Coelophysis + Megapnosaurus, as defined in Tykoski and Rowe (2004), then Segisauridae may be used for its sister clade, including Segisaurus, "Megapnosaurus" kayentakatae and perhaps Camposaurus. On this site they are grouped as Segisaurinae within Coelophysidae.
Ex-coelophysids- Huene (1914) included Saltopus as one of his original podokesaurids, placing it ancestral to Halticosaurus. This placement was common, though current research indicates it is most likely a more basal dinosauriform (Langer, 2004). Tanystropheus is another original podokesaurid of Huene's, but is now known to be a protorosaur (Wild, 1973). Several species (T. bauri, T. longicollis and T. willistoni) are now recognized as coelophysids however. Halticosaurus was assigned to Podokesauridae (Huene, 1914) by many authors and indeed seems to be coelophysoid, though its short cervical centra exclude it from Coelophysidae as used here. Procompsognathus was soon placed in Podokesauridae by Huene (1914), which has been supported by some recent analyses (Ezcurra and Novas, 2006) and rejected by others (Allen, 2004). Procerosaurus was assigned to Podokesauridae by Huene (1920), but is actually a junior synonym of Tanystropheus (Wild, 1973). Spinosuchus was first described as a Coelophysis specimen (Case, 1922), and was later referred to Podokesauridae once it was placed in its own genus by Huene (1932). Nesbitt et al. (2007) have most recently assigned it to Archosauriformes incertae sedis. Avipes was first described as a podokesaurid (Huene, 1932), but recent studies have been unable to place it more definitively than Avemetatarsalia (Rauhut and Hungerbuhler, 2000). Lukousaurus was originally described as a coelurosaur related to Podokesaurus, Procompsognathus and Saltopus (Young, 1948), and explicitly placed in Podokesauridae by Romer (1956). It has not been recently restudied, but lacks several coelophysoid apomorphies and is more likely crurotarsan. Velocipes was placed in Podokesauridae by Romer (1956), yet like Avipes, recent studies have been unable to place it more definitively than Avemetatarsalia (Rauhut and Hungerbuhler, 2000). Dolichosuchus was similarly placed in Podokesauridae by Romer (1956), but Welles (1984) notes it is almost identical to Liliensternus, so it is possibly just outside Coelophysidae. Reig (1963) assigned Trialestes (as Triassolestes) to Podokesauridae. Romer (1972) properly recognized it as a crurotarsan, though it does have several dinosaur-like characters. Russell (1984) assigned Dilophosaurus to the Podokesauridae, though following Holtz (1994), it is now considered just basal to Coelophysidae. Welles (1984) placed Sarcosaurus in Coelophysidae without comment, while Madsen and Welles (2000) placed it in Podokesauridae, but it appears to represent a more basal coelophysoid (Tykoski, 2005). Protoavis was originally believed to be a juvenile Coelophysis (Chatterjee, 1986), and while some material is coelophysoid (Nesbitt et al., 2007), none can be assigned to Coelophysidae itself with certainty. (Chatterjee (1987) assigned his new genus Walkeria to Podokesauridae, but it has since been renamed Alwalkeria and placed more basally as a basal eusaurichian (Langer, 2004). Elaphrosaurus was assigned to Coelophysidae by a couple authors (Paul, 1988; Novas, 1992) before it was realized to be a ceratosaur sensu lato (Holtz, 1994), which has been the result of every published cladistic analysis. Liliensternus is often assigned to Coelophysidae (e.g. Paul, 1988) and was often viewed as a podokesaurid when still classified as a species of Halticosaurus, yet based on the present definition of Coelophysidae is just barely excluded from that clade. "Comanchesaurus" was listed as a podokesaurid by Hunt and Lucas (1989), though it more recently could not be definitely placed closer to Coelophysis than Liliensternus (Nesbitt et al., 2007). Sullivan and Lucas (1999) believed their new taxon Eucoelophysis to be most closely related to Coelophysis and Megapnosaurus, but Ezcurra (2006) has since determined it to be a more basal dinosauriform. Rauhut (2003) found Shuvosaurus to fall out as a coelophysid in his analysis, sister to Megapnosaurus, but the description of Effigia (Nesbitt and Norell, 2006) led to its recognition as a crurotarsan.
Defining Coelophysidae- Holtz (1994) first defined Coelophysidae, as the most exclusive clade containing Coelophysis bauri, Syntarsus rhodesiensis and S. kayentakatae. This matches its use in recent papers such as Tykoski (2005) and Ezcurra and Novas (2006). Two other definitions have been proposed for Coelophysidae, neither one ideal. Sereno (1998) used Procompsognathus and Coelophysis as internal specifiers. Yet Procompsognathus may not be dinosaurian (Allen, 2004), in which case all of Dinosauria would be coelophysids according to his definition. Even when Procompsognathus is found to be a coelophysoid, it usually has an uncertain position within the clade. Tykoski (2005) found it could fall it anywhere in the Liliensternus + Coelophysis clade. Ezcurra and Novas (2006) found it only slightly more constrained, within the Zupaysaurus + Coelophysis clade. Under this definition, Coelophysidae has an extremely uncertain content, which may include Liliensternus, Zupaysaurus, Segisaurus, etc. or may not. Tykoski and Rowe (2004) proposed a definition which used Coelophysis and Megapnosaurus rhodesiensis as internal specifiers. This is easy to apply to most analyses, where the two emerge as sister taxa. However, this would exclude not only Liliensternus, Zupaysaurus and Lophostropheus from Coelophysidae, but also Segisaurus and "Megapnosaurus" kayentakatae. This is far more exclusive than most concepts of Coelophysidae (or Podokesauridae) have been. Also, the synapomorphies for Coelophysis + Megapnosaurus are all cranial, making the assignment of postcrania to Coelophysidae impossible.
References- Huene, 1914a. Beiträge zur geschichte der Archosaurier [Contribution to the history of the archosaurs]. Geologie und Paläontologie Abhandlungen. 13(7), 1-56.
Huene, 1914b. Das natürliche System der Saurischia [The systematics of the Saurischia]. Centralblatt für Mineralogie, Geologie und Paläontologie. 1914,154-158.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien [Phylogenetic results of some investigations of Triassic reptiles]. Zeitschrift für Induktive Abstammungsund Vererbungslehre. 24, 159-163.
Case, 1922. New reptiles and stegocephalians from the Upper Triassic of Western Texas. Carnegie Institution of Washington Publication. 321, 1-84.
Nopcsa, 1923. Die Familien der Reptilien [The families of reptiles]. Forschritte der Geologie und Palaeontologie. 2, 1-210.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Huene, 1929. Kurze Übersicht über die Saurischia und ihre natürlichen Zusammenhänge [A brief survey of the Saurischia and their natural context]. Paläontologische Zeitschrift. 11, 269-273.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monog. Geol. Pal. 1, pts 1 and 2, V111 + 361 pp.
Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of Arizona. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 24 39-56, 8 figs., 2 pls.
Young, 1948. On two new saurischians from Lufeng, China. Bulletin of the Geological Society of China. 28(1-2), 78-90.
Romer, 1956. Osteology of the Reptiles, University of Chicago Press. 1-772.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press, Chicago. 1-468.
Raath, 1969. A new coelurosaurian dinosaur from the Forest Sandstone of Rhodesia. Arnoldia (Rhodesia). 4(28), 1-254.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 1-87.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse) . Schweizerische Palaontologische Abhandlungen. 95, 1-162.
Barsbold, 1977. O evolutsiy chishcheich dinosavrov [On the evolution of carnivorous dinosaurs]. Trudy - Sovmestnaya Sovetsko-Mongol'skaya Paleontologicheskaya Ekspeditsiya. 4, 48-56.
Russell, 1984. A check list of the families and genera of North American dinosaurs. Syllogeus. 53, 1-35.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Chatterjee, 1986. The Late Triassic Dockum vertebrates: Their stratographic and paleobiogeographic significance. In Padian (ed.). The beginning of the age of dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge University Press. 378 pp.
Chatterjee, 1987. A new theropod dinosaur from India with remarks on the Gondwana-Laurasia connection in the Late Triassic. Gondwana 6: Stratigraphy, Sedimentology and Paleontology, G. D. McKenzie (ed.), Geophysical Monographs. 41, 183-189.
Carroll, 1988. Vertebrate Paleontology and Evolution. W.H. Freeman and Company.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Hunt and Lucas, 1989. Late Triassic vertebrate localities in New Mexico. pp. 72-101. in Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque.
Novas, 1991. Relaciones filogeneticas de los dinosaurios teropodos ceratosaurios [Phylogenetic relationships of ceratosaurian theropod dinosaurs]. 28(3-4), 401.
Novas, 1992. La evolucion de los dinosaurios carnivoros [The evolution of carnivorous dinosaurs]. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico: Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez", Cuenca, Argentina. 126-163.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences. 25, 435-489.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 210(1), 41-83.
Sullivan and Lucas, 1999. Eucoelophysis baldwini, a new theropod dinosaur from the Upper Triassic of New Mexico, and the status of the original types of Coelophysis. Journal of Vertebrate Paleontology 19(1): 81-90.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised osteology. Miscellaneous Publication 00-2 Utah Geological Survey, 80 pages.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia. 15, 75-88.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria:Theropoda) from the Late Cretaceous of Madagascar. Journal of Vertebrate Palaeontology. 22, 510–534.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP 24(3).
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas. 28 (4),649-684.
Nesbitt and Norell, 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B. 273, 1045-1048.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

unnamed Coelophysidae (Cope, 1887)
Late Norian, Late Triassic
Petrified Forest Member of the Chinle Formation, New Mexico, US
Material- ?(AMNH 2702; syntype of Coelurus longicollis) mid caudal vertebra (51 mm) (Cope, 1887a)
?(AMNH 2703) manual ungual (Cope, 1887)
(AMNH 2706) pubis (228 mm) (Cope, 1887)
?(AMNH 2707) manual phalanx I-1 (43 mm) (Cope, 1887)
(AMNH 2708; not lectotype of Coelophysis bauri, contra Welles, 1984) ilium (~155 mm) (Cope, 1887b)
?(AMNH 2715; syntype of Coelurus longicollis) dorsal centrum (42 mm) (Cope, 1887a)
?(AMNH 2716) ischium(?) (Cope, 1887)
(AMNH 2717; syntype of Coelurus bauri) third or fourth cervical centrum (53 mm) (Cope, 1887a)
(AMNH 2718) distal ischium (Cope, 1887)
?(AMNH 2719) proximal ischium(?) (Cope, 1887)
(AMNH 2720) partial fourth or fifth cervical vertebra (Cope, 1887)
(AMNH 2722; lectotype of Coelophysis bauri) sacrum (20, 19, 19, 16, ? mm) (Cope, 1887a)
?(AMNH 2723) dorsal centrum (30 mm) (Cope, 1887)
(AMNH 2724) proximal pubis (Cope, 1887)
?(AMNH 2727; syntype of Tanystropheus willistoni) distal caudal centrum (Cope, 1887b)
?(AMNH 2728) distal metatarsal IV(?) (Huene, 1915)
?(AMNH 2729) dorsal transverse process (Huene, 1915)
?(AMNH 2730) distal metatarsal III (Huene, 1906)
?(AMNH 2731) proximal pubis (Huene, 1915)
?(AMNH 2732) five vertebral fragments (Padian, 1986)
?(AMNH 2734) incomplete mid caudal vertebra (Huene, 1915)
?(AMNH 2735) partial distal caudal centrum (Cope, 1887)
?(AMNH 2736) partial cervical neural arch (Huene, 1915)
?(AMNH 2737) proximal humerus (Huene, 1915)
?(AMNH 2738) ischial fragments (Huene, 1915)
?(AMNH 2739) dorsal centrum (Huene, 1915)
?(AMNH 2740) distal metatarsal III (Huene, 1915)
?(AMNH 2742) bone fragments (Padian, 1986)
?(AMNH 2743) vertebra, six fragments (Padian, 1986)
?(AMNH 2744) partial proximal caudal vertebra (Huene, 1906)
?(AMNH 2745) proximal fibula (Huene, 1915)
?(AMNH 2746) five caudal vertebra fragments (Padian, 1986)
?(AMNH 2747) pedal fragments (Padian, 1986)
?(AMNH 2748) vertebral fragments (Padian, 1986)
?(AMNH 2749) dorsal centrum (Huene, 1915)
?(AMNH 2750) anterior sacrum (Huene, 1915)
?(AMNH 2751) posterior cervical vertebra (Huene, 1915)
?(AMNH 2752) partial anterior dorsal vertebra (Huene, 1915)
?(AMNH 2753) vertebral and bone fragments (Padian, 1986)
Comments- This material was collected with the lectotypes of Coelophysis bauri, C. longicollis, C. willistoni and the holotype of Longosaurus. It has been referred to various combinations of these species by different authors (Cope, 1887, 1887; Huene, 1906, 1915; Welles, 1984; Padian, 1986). Most recently, it was viewed as generically indeterminate (along with the types of the above species), and Coelophysis was assigned a neotype from a different locality (Hunt and Lucas, 1991; ICZN, 1996). Sullivan and Lucas (1999) erected a new species of coelophysid from the Coelophysis lectotype locality, Eucoelophysis baldwini, and referred AMNH 2706 to it. In addition, they felt the rest of the material listed above was probably referrable to Eucoelophysis, but could not definitively assign it due to its undiagnostic nature. Nesbitt et al. (2005, 2007) and Ezcurra (2006) determined Eucoelophysis was not a theropod, but a more basal dinosauriform. Nesbitt et al. (2007) reexamined the specimens and found that the cervical vertebrae (AMNH 2717, 2720), sacrum (AMNH 2722) and pelvic elements (AMNH 2705, 2706, 2708, 2722) could be referred to coelophysoids based on- dual pairs of cervical pleurocoels, pubic obturator foramen, well developed supraacetabular crest that arcs ventrally at its lateral margin; a squared-off distal portion of the postacetabular process; a deep brevis fossa where the lateral ridge originates near the supraacetabular crest; flattened dorsal margin of the iliac blade; fully perforated acetabulum. It's probable only one species of coelophysid is represented, possibly congeneric or conspecific with C. bauri. The dorsal vertebrae and most of the limb elements were undiagnostic within Archosauria, the supposed proximal tibia (AMNH 2721) is referrable to Dromomeron romeri (Nesbitt et al., 2009; actually a distal femur), and a distal femur (AMNH 2725) is referrable to a shuvosaurid.
References- Cope, 1887. The dinosaurian genus Coelurus. American Naturalist. 21, 367-369.
Cope, 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls. i, ii.
Huene, 1906. Ueber die Dinosaurier der Aussereuropaischen Trias. Geologische und Paläontologische Abhandlungen. 12, 99-156.
Huene, 1915. On reptiles of the New Mexican Trias in the Cope collection. Bulletin American Museum of Natural History. 34, 485-507.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic dinosaur from New Mexico (USA). Paläontol. Z. 65 p. 191-198.
International Commision on Zoological Nomenclature, 1996. Opinion 1842. Coelurus bauri Cope, 1887 (currently Coelophysis bauri; Reptilia, Saurischia): lectotype replaced by a neotype. Bulletin of Zoological Nomenclature. 53, 142-144.
Sullivan, Lucas, Heckert and Hunt, 1996. The type locality of Coelophysis, a Late Triassic dinosaur from north-central New Mexico (USA). Palaeontologische Zeitschrift. 70(1/2):245-255.
Sullivan and Lucas, 1999. Eucoelophysis baldwini, a new theropod dinosaur from the Upper Triassic of New Mexico, and the status of the original types of Coelophysis. Journal of Vertebrate Paleontology 19(1): 81-90.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. JVP. 25(3) 96A.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas. 28 (4),649-684.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.
Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. Journal of Vertebrate Paleontology. 29(2), 498-516.

Coelophysidae indet. (Heckert, Zeigler, Lucas, Rinehart and Harris, 2000)
Late Norian, Late Triassic
Petrified Forest Member of Chinle Formation, New Mexico
Material- (NMMNH P-29168) partial tibiotarsus fused to fibula
References- Heckert, Zeigler, Lucas, Rinehart and Harris, 2000. Preliminary description of coelophysoids (Dinosauria: Theropoda) from the Upper Triassic (Revueltian: early-mid Norian) Snyder quarry, north-central New Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 27-32.
Heckert, Zeigler, Lucas and Rinehart, 2003. Coelophysids (Dinosauria: Theropoda) from the Upper Triassic (Revueltian) Snyder quarry. New Mexico Museum of Natural History and Science Bulletin. 24, 127-132.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.
Spielmann, Lucas, Rinehart, Hunt, Heckert and Sullivan, 2007. Oldest records of the Late Triassic theropod dinosaur Coelophysis bauri. New Mexico Museum of Natural History and Science Bulletin. 41, 384-401.

undescribed coelophysid (Taquet, 1984)
Toarcian, Early Jurassic
Wazzant Formation, Morocco
Material- (MNHN coll.) cervical vertebra, fibula, partial astragalus, calcaneum, distal tarsal III, metatarsal I, phalanx I-1, ungual I, metatarsal II, phalanx II-1, phalanx II-2, ungual II, metatarsal III (125 mm), phalanx III-1, proximal phalanx III-2, metatarsal IV, phalanx IV-1, phalanx IV-2, metatarsal V
Comments- This is provisionally assigned to Coelophysidae based on the apparent fusion between distal tarsal III and metatarsal III. Allain et al. (2007) note it is currently under preparation at the MNHN, has long cervical centra and lacks a medial fibular sulcus.
References- Jenny, Jenny-Deshusses, Le Marrec and Taquet, 1980. Decouverte d'ossements de Dinosauriens dans le Jurassique inferieur (Toarcien) du Haut-Atlans central (Maroc). Comptes Rendus Academie des Sciences. 290 839-842.
Taquet, 1984. Two new Jurassic specimens of coelurosaurs (Dinosauria). in Hecht, Ostrom, Viohl and Wellnhofer (eds). The Beginnings of Birds: Proceedings of the International Archaeopteryx Conference, Eichstaett. 229-232.
Allain, Tykoski, Aquesbi, Jalil, Monbaron, Russell and Taquet, 2007. An abelisauroid (Dinosauria: Theropoda) from the Early Jurassic of the High Atlas Mountains, Morocco, and the radiation of ceratosaurs. Journal of Vertebrate Paleontology. 27(3), 610-624.

unnamed coelophysid (Fraser and Padian, 1995)
Norian?, Late Triassic
Pant-y-ffynnon fissure filling, Wales
Material- (BMNH PV RU 76/1, 77/1) posterior dorsal vertebrae, partial sacrum, ilium (52 mm), incomplete pubis, partial ischium, incomplete femur
Comments- Rauhut and Hungerbuhler (2000) state it is extremely similar to Megapnosaurus rhodesiensis in all features, although it is also very similar to Procompsognathus and may prove to be referrable to that genus.
References- Fraser and Padian, 1995. Possible dinosaur remains from Britain and the diagnosis of the Dinosauria. J.Vert.Paleont. 15(3): 30A.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.

Shake-N-Bake coelophysid (Tykoski, 1997)
Hettangian, Early Jurassic
Kayenta Formation, Arizona, US
Material- (MCZ 8817, 9442-9469; TMM 43689) (at least fifteen individuals, adults) several hundred specimens including several dorsal vertebrae, several sacra, many caudal vertebrae, several ilia, fifteen proximal femora, several distal femora, six distal tibiotarsi (9.4, 11.1, 12.6, 12.9, 13 mm transversely), including...
(MCZ 8817b) dorsal vertebra, partial sacrum, partial pelvis
(MCZ 8817c) two partial mid cervical vertebrae
(MCZ 8817d) incomplete posterior cervical vertebra
(MCZ 8817e) proximal caudal vertebra
(MCZ 8817f) partial tooth, partial caudal centrum
(MCZ 8817g) anterior dorsal vertebra
(MCZ 8817h) incomplete anterior dorsal vertebra
(MCZ 8817i) partial sacrum
(MCZ 8817j) proximal femur
(MCZ 8817k) (robust) proximal femur
(MCZ 8817l) (gracile) proximal femur
(MCZ 8817m) distal tibiotarsus
(MCZ 8817n) distal tibiotarsus, distal fibula
(MCZ 8817o) distal tibiotarsus, distal fibula
(MCZ 8817p) distal tibiotarsus
(MCZ 8817q) distal tibia, partial astragalus
(MCZ 8817r) partial scapulocoracoid
(MCZ 8817t) partial scapulocoracoid
(MCZ 9442; = 8817a) (adult) sacrum, partial ilia, proximal pubes, proximal ischia
(MCZ 9463; = 8817m-p?) (adult) distal tibiotarsus
(TMM 43689-4) (adult) proximal tarsometatarsus
Comments- This new taxon is based on specimens formerly thought to be juvenile "Megapnosaurus" kayentakatae by Rowe (1989). Initially catelogued at MCZ 8817 (Tykoski, 1998), at least some have since been recatalogued as MCZ 9442-9469 and TMM 43689 (Tykoski, 2005). Tykoski (1998) used MCZ 8817a-s as preliminary labels for some specimens, but it is unknown which numbers most of these specimens are now catalogued under.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. J. Vertebr. Paleontol. 9 p. 125-136.
Tykoski, 1997. A new ceratosaurid theropod from the Early Jurassic Kayenta Formation of Northern Arizona. JVP 17(3) 81A-82A
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications for ceratosaurid phylogeny. Unpublished Masters Thesis,University of Texas at Austin, 217 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.

unnamed coelophysid (Lucas and Heckert, 2001)
Hettangian, Early Jurassic
Moenave Formation, Arizona, US
Material- (MNA V1968) several dorsal ribs or gastralia, incomplete ilium, proximal pubis, proximal femur, incomplete ischium, distal ischium
(MNA V2588) five incomplete proximal caudal centra (34-37 mm), ilium (196 mm), incomplete pubis, incomplete ischia (172 mm)
Comments- Pubes fuse to ischia posterior to acetabular midpoint, unlike Coelophysis or Megapnosaurus species. Position ischia fuse to ilia on antitrochanter more similar to "Megapnosaurus" kayentakatae than Coelophysis. Medial bowing of femur more similar to M. rhodesiensis than "M." kayentakatae.
References- Morales, 1994. First dinosaur body fossils from the Lower Jurassic Dinosaur Canyon Member, Moenave Formation of Northeastern Arizona. JVP. 14(3), 39A.
Lucas and Heckert, 2001. Theropod dinosaurs and the Early Jurassic age of the Moenave Formation, Arizona-Utah, USA. Neues Jahrbuch fur Geologie und Palaontologie Monatshefte 2001(7): 435-448.

unnamed coelophysid (Irmis, 2004)
Hettangian-Pliensbachian, Early Jurassic
Lower Lufeng Formation, Yunnan, China
Material- (FMNH CUP 2089) vertebra?, distal humerus, proximal radius, proximal ulna, metacarpal fragments, manual phalangeal fragments
....(FMNH CUP 2090) distal tarsal II, distal tarsal III, proximal metatarsal II, proximal metatarsal III
Comments- Simmons (1965) identified FMNH CUP 2089 as a podokesaurid distal femur, proximal tibia and proximal fibula. Irmis (2004) referred it to Megapnosaurus due to the fusion of metatarsals II and III, but the character is present in Coelophysis bauri and the Shake-n-Bake coelophysid too (Tykoski, 2005).
References- Simmons, 1965. The non-therapsid reptiles of the Lufeng Basin, Yunnan, China. Fieldiana: Geology 15:1–93.
Irmis, 2004. First report of Megapnosaurus (Theropoda: Coelophysoidea) from China. PaleoBios. 24(3), 11-18.

undescribed Coelophysidae (UCMP online)
Early Norian, Late Triassic
Mesa Redondo Member of the Chinle Formation, Arizona, US
Material- (UCMP 25968) ilium
(UCMP 25971) pubis
(UCMP 25974) ilium
(UCMP 25978) ilium
(UCMP 25982) ilium
(UCMP 25990) ischium
(UCMP 25993) ischium
(UCMP 25994) ischium
(UCMP 27982) ilium
Comments- This material was collected in 1934 and is referred to Coelophysis on the UCMP collections website.
Reference- U.C.M.P. Database. 2005. UCMP collections database. University of California Museum of Paleontology.

undescribed Coelophysidae (UCMP online)
Late Triassic
Dockum Group, Texas, US
Material- (UCMP 65737) cervical vertebral fragments
(UCMP 65773) dorsal vertebra
(UCMP 65774) caudal vertebra
(UCMP 65775)
Reference- U.C.M.P. Database. 2005. UCMP collections database. University of California Museum of Paleontology.

Coelophysis? willistoni (Cope, 1887) Cope, 1889
= Tanystophaeus willistoni Cope, 1887
Late Norian, Late Triassic
Petrified Forest Member of the Chinle Formation, New Mexico, US
Lectotype- (AMNH 2726) partial ilium (160 mm)
Comments- This was originally named based on the lectotype and an unassociated distal caudal centrum. Further material from the same locality was later included in the hypodigm, but the lack of association prevents referring anything to this taxon except the lectotype ilium. See the discussion of "unnamed Coelophysidae (Cope, 1887)" above for more details. The lectotype is probably indeterminate at the level of Coelophysidae. Although coelophysid synapomorphies aren't preserved in the holotype, the coelophysoid-like angled pubic articular surface of the ilium and association with coelophysid elements indicate the specimen was probably a coelophysid itself.
References- Cope, 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls. i, ii.
Cope, 1889. On a new genus of Triassic Dinosauria. Amer. Naturalist xxiii p. 626.

Kayentavenator Gay, 2010
K. elysiae Gay, 2010
Hettangian, Early Jurassic
Silty Facies Member of the Kayenta Formation, Arizona, US
Holotype- (UCMP V128659) (juvenile) six proximal caudal centra, three centra, two partial neural arches, fragmentary ilium, proximal pubes, pubic shaft fragments, incomplete femora, proximal tibiae, proximal fibula, fragments
Diagnosis (modified from Gay, 2010) short anteriorly projected pubic peduncle on the ilium (possibly caused by breakage); accessory medial femoral condyle (possibly the ectocondylar tuber misidentified due to switching left and right femora); mediodistal femoral crest longer than half of shaft length (exact position and thus homlogy uncertain).
Other diagnoses- Gay lists several characters in the diagnosis which are meant to distinguish it from other theropods, including kayentakatae. Gay's description of Kayentavenator's femoral condyles is confusing as the "accessory condyle" is said to project from the medial condyle, yet the only accessory condyle in theropods including birds (which Gay states the accessory condyle resembles) is the ectocondylar tuber which is associated with the lateral condyle. Unfortunately, this supposedly unique morphology is not illustrated, with the femur only photographed in anterior view. Since the distal femur is separated from the proximal end in at least one element (and presumably the other, as the total length of both is said to be difficult to determine), it seems at least possible Gay confused the right and left distal femora. This would give them standard theropod ectocondylar tubers instead of apomorphically lacking the tuber and having a unique medial accessory condyle. The acetabular shape is described as arching "upward under the acetabular rim, making it taller than it is wide." However, it is clear from the illustration and Tykoski's photo that the supracetabular crest is broken. If it was pendent as in other coelophysoids, the apparent acetabular depth would decrease. If the below speculation about the pubic peduncle being broken in Gay's illustration is correct, this would also add to the acetabulum's length once corrected for. The greater trochanter and femoral head are stated to be fused, which seems to be an unorthodox way of saying there is no concavity between them. Gay notes in the description Coelophysis shares this morphology, and this is true for all other coelophysoids as well. Gay describes a mediodistal crest on the femur extending from the medial condyle for at least half the length of the element. While this might be assumed to be the medial epicondyle on the anteromedial edge common in non-tetanurines (but always less than a third of femoral length), kayentakatae also has a longer (40% of femoral length) crest extending from the medial condyle on the posterior shaft. Of course if the distal femora have been switched as proposed above, the crest would become lateral instead. A posterior lateral crest is present in kayentakatae's holotype, but this is intermediate in length between the medial epicondyle and postromedial crest. These posterior crests are undeveloped or poorly developed in Dilophosaurus, Liliensternus, Coelophysis and Segisaurus, though seem present in at least some Megapnosaurus. The distal femur is photographed in anterior view and shows no evidence of a medial epicondyle (though the dark coloring in this area could indicate the bone surface is broken off), while any posterior crest is of course unobservable. The transverse groove on the proximal femoral head surface is said to be unique among theropods, but is polymorphic in Coelophysis (e.g. NMMNH P-29046 and P-54620, UCMP 129618), so could be expected in some kayentakatae individuals as well. The caudal centra are said to be highly constricted, with the description further specifying "minimum width of approximately 4mm, with an articular surface diameter of 17mm (Figure1)." Based on the scale in that figure, the articular surface is indeed close to 17 mm, but the minimum central height is 11 mm instead of 4 mm. While kayentakatae caudals have not been illustrated in lateral view, these proportions are similar to other coelophysoids like Megapnosaurus and Liliensternus. A few additional characters are listed as being specifically distinct from kayentakatae. The anterior trochanter is said to be placed more medially, but this is not true. It should be noted that only robust femora are otherwise known for kayentakatae, which makes comparison of minor details questionable. The trochanter of Kayentavenator is actually more laterally placed than gracile individuals of Dilophosaurus and Megapnosaurus, but comparable to Liliensternus. The "groove in ventral surface of femoral head" is presumedly a typo for the groove in the proximal surface, which is dealt with above. The "spike on medial surface of tibia" is a typo for the lateral fibular crest, as indicated by Gay using the same character with 'lateral' substituted for 'medial' to distinguish Kayentavenator from Coelophysis, Megapnosaurus and Dilophosaurus. The crest of course is not a spike, and is stated to be large in kayentakatae as well. Thus the supposed diagnostic characters are all problematic. The accessory femoral condyle and acetabular shape may be misinterpreted, the distal femoral crest is unique as described but impossible to homologize, the caudal proportions and "fused" greater trochanter are normal for coelophysoids, and the transverse femoral head groove is prone to individual variation.
Comments- UCMP V128659 was discovered in 1982 and referred to Syntarsus kayentakatae by Rowe (1989), as a subadult gracile individual. Tykoski (1998) did not examine it for his redescription of the species, but later (2005) examined it for his PhD thesis and considered it to be "probably referrable to "Syntarsus" kayentakatae" without discussion. Gay (2010) described the specimen as the new taxon Kayentavenator elysiae.
Kayentavenator a tetanurine? Based on a small phylogenetic analysis, Gay placed Kayentavenator in Tetanurae but outside Avetheropoda. This was based on several characters. The pubic articulation of the ilium is also larger than the ischial articulation in kayentakatae, and by a larger amount than Kayentavenator. The pubic peduncle's distal surface is also longer than wide in ceratosaurs and coelophysids like Megapnosaurus. The cnemial process arises from the lateral surface of the tibia in almost all theropods including kayentakatae. The trochanteric shelf is absent in all gracile and juvenile ceratosaurs and coelophysids, so cannot be used to place the juvenile Kayentavenator holotype in Tetanurae. Finally, the anterodistal femoral fossa is said to be non-elliptical in shape, which refers to a character originally used by Perez-Moreno et al. (1993). Ironically, in Perez-Moreno et al.'s analysis, the avetheropods were coded as having an elliptical fossa unlike Gay's analysis. In truth, avetheropods do not have fossae that are more or less oval than that of more basal theropods. While the fossa is poorly developed in Coelophysis, it is illustrated by Rowe in kayentakatae (as being non-elliptical due to its flat medial edge, for what it's worth) and is stated to be distinct in Segisaurus as well. Gay also lists a feature in the description that is supposedly diagnostic of tetanurines- a pronounced sheet of bone projecting from the medial surface of the tibia, referring to Naish's (1999) description of BMNH R9385. Yet this must be a mistake as the feature Naish describes is the fibular crest on the lateral surface. However, Segisaurus has a prominent fibular crest comparable to tetanurines', while kayantakatae's is also described as large. There are therefore no characters placing Kayentavenator in Tetanurae.
Kayentavenator not a coelophysid? Gay states Kayentavenator "lacks a crista tibiofibularis and its associated groove, which are present in all coelophysoids and Dilophosaurus." Yet coelophysoids do not have a tibiofibular crest, the structure labeled as such by Rowe in kayantakatae being the ectocondylar tuber present in almost all theropods. Young kayentakatae and Dilophosaurus specimens lack the deep groove lateral to the ectocondylar tuber, so its absence in the juvenile Kayentavenator specimen (confirmed by Tykoski, 2005) is expected. Kayentavenator is coded differently than "Coelophysisidae" [sic] in Gay's matrix for several additional characters. The caudal vertebrae are coded as having pleurocoels in the neural arch, which is not possible since pleurocoels are by definition a feature of vertebral centra. Furthermore, Gay states the position of the two partial preserved neural arches is impossible to ascertain. The pneumatic fossae are stated to face anteriorly on each side of the neural arch, indicating they may be anterior peduncular fossae as in Coelophysis cervicals, or even anterior infradiapophyseal fossae which all theropod presacrals possess. Since the neural arches are so fragmentary they cannot even be placed in the vertebral column (they are assumed by Gay to be from the posterior region only because the other remains are from the pelvis and hindlimb), they could even be backwards and merely exhibit posterior peduncular fossae as in all coelophysid cervicals including those of kayentakatae. Again, this supposedly unique feature is not illustrated, making evaluation difficult. The brevis fossa is coded as being deep unlike coelophysoids, but coelophysoids including Megapnosaurus have deep brevis fossae. Coelophysids are oddly coded as lacking a supracetabular crest, which is untrue. They are also incorrectly coded as having an acetabular height only a third or less of the acetabular length, which is not true of any theropod (e.g. the ratio in kayentakatae is 88%). For the character "Pubic peduncle of ilium depth: 0, extends ventrally to the same level as ischiadic peduncle; 1, extends more ventrally than ischiadic peduncle.", coelophysids are coded as having nonexistant state 2 unlike Kayentavenator's state 0. In actuality coelophysids including kayentakatae have pubic peduncles extending ventral to their ischial peduncle. This is also true in Kayentavenator based on the stereophotograph in Tykoski's (2005) thesis, which does not agree with Gay's drawing. The photo also shows a complete articular surface on the pubic peduncle, while no obvious anteroventral corner to the process exists in Gay's illustration. Perhaps the peduncle was broken off during Gay's examination? The obturator foramen is coded as open in Kayentavenator, despite Gay illustrating the ventral edge as closed but broken and stating the ventral margin was missing. Oddly, Kayentavenator is coded as having a pubic fenestra while coelophysids are not, despite the fact the latter are the theropods best known for having pubic fenestrae. Gay codes Kayentavenator as having a more propubic pelvis (~30 degrees from horizontal) than coelophysids (~45 degrees). This would be based off the angle of the pubic peduncle's articular surface, but as noted above, the preservation of this surface in Gay's illustration is in doubt. The femoral head is coded as being subequally long and deep (in anterior/posterior view) while coelophysids' are coded as proximodistally elongate. However, the transverse width (from medial edge of femoral head to medial edge of shaft) is only 68% of the proximodistal height of the head, which is close to that in the kayentakatae holotype (63%). Since Coelophysis varies between 43 and 62%, a difference of 5% seems within plausible individual variation in kayentakatae. The anterior trochanter is aliform (as confirmed by Tykoski, 2005) while coelophysids' were incorrectly coded as absent. In actuality, many gracile coelophysoids (e.g. Dilophosaurus, Megapnosaurus) have aliform anterior trochanters as well. Gay codes coelophysids as having an anterior trochanter (contra the previous character) which does not extend proximally past the femoral head's ventral margin unlike Kayentavenator, but coelophysids' anterior trochanters do in fact extend past the femoral head's margin (e.g. kayentakatae- Rowe, 1989). Finally, the proximomedial fibular sulcus is coded as absent in coelophysids unlike Kayentavenator, but this feature is present in all adult coelophysids and was even made famous by kayentakatae. In all, the characters which supposedly differ from coelophysids are miscodings or based on questionable morphologies (perhaps switched distal femora, possibly broken pubic peduncle, uncertain neural arch position). Supporting the placement of Kayentavenator in the Coelophysoidea is the presence of a divided articular facet on the pubic peduncle of the ilium, as illustrated by Tykoski.
Kayentavenator an individual of kayentakatae? Comparing Kayentavenator to other coelophysoids is made difficult not only by the poor preservation and juvenile status of the former, but also the wanting description and figures, as well as the general lack of postcranial characters in coelophysid diagnoses. The one complete caudal centrum of Kayentavenator lacks a ventral median groove, unlike at least some of kayantakatae's centra. Yet this varies within the tail of many theropods like Eustreptospondylus, so is probably unimportant. The pubic peduncle being longer than wide is more similar to Megapnosaurus than Dilophosaurus and Liliensternus, as is the narrow and ventrally pointed ischial peduncle. The femoral head is more elongate than other coelophysoids (including Halticosaurus- contra Gay), but this shows individual variation that could be accommodated by kayentakatae as noted above. If Gay's correct about the anterodistal femoral fossa, this is like Megapnosaurus, Segisaurus and kayentakatae but unlike Coelophysis, Liliensternus and Dilophosaurus. The fibular crest is larger in Segisaurus than in Coelophysis, which is in turn larger than in Dilophosaurus. Based on Gay's description, Kayentavenator is more similar to coelophysids in this respect, but without figures it's difficult to determine. Similarly, any comparisons of vertebral fossae or distal femoral ridges are hindered by their unknown homology due to a lack of description and figures. Based on published evidence, Kayentavenator seems to be a coelophysid. Past that, it's difficult to tell. There's nothing verified that is more similar to kayentakatae than to other coelophysids, so I don't think it should be referred to that species. There are already at least two Kayenta coelophysids after all (kayentakatae and the Shake-n-Bake taxon which is too small and fused to belong to Kayentavenator). There are a few supposed diagnostic characters that have escaped definite rejection (short anteriorly projected pubic peduncle on the ilium; accessory medial femoral condyle; ambiguous mediodistal femoral crest longer than half of shaft length), but I'm hesitant to believe these are real based on the lack of appropriate femur illustrations, differences from the ilium's photo in Tykoski's thesis, and generally large amount of errors present in the paper. Further analysis may vindicate Gay or may identify features shared only with kayentakatae. At the moment, whether one makes Kayentavenator a nomen dubium depends on how much one trusts Gay's description.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9, 125-136.
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications for ceratosaurid phylogeny. Unpublished Masters Thesis, University of Texas at Austin, 217 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Gay, 2010. Notes on Early Mesozoic theropods. Lulu Press. 44 pp.

Longosaurus Welles, 1984
L. longicollis Welles 1984
Late Norian, Late Triassic
Petrified Forest Member of the Chinle Formation, New Mexico, US
Holotype- (AMNH 2705) ilium (189 mm)
Comments- Welles (1984) intended to separate Coelophysis longicollis from Coelophysis bauri at the genus level by naming this genus, but accidentally based it on a different specimen than the lectotype of Tanystropheus longicollis. Thus these taxa have different type specimens and are not objective synonyms. In addition, Welles referred at least some material Huene (1912) referred to Coelophysis longicollis to Longosaurus longicollis (AMNH 2701, 2703, 2704, 2707). See the discussion of "unnamed Coelophysidae (Cope, 1887)" above for more details. The lectotype is probably indeterminate at the level of Coelophysidae. The concave posterior postacetabular process margin indicates it is a coelophysid however.
Reference- Cope, 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls. i, ii.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica 185A, 85-180.

Podokesaurus Talbot, 1911
P. holyokensis Talbot, 1911
= Coelophysis holyokensis (Talbot, 1911)
Pliensbachian-Toarcian, Early Jurassic
Portland Formation?, Massachusetts, US
Holotype- (destroyed) (.89 m; .92 kg) cranial elements, four cervical vertebrae (12-13 mm), cervical ribs, thirteen dorsal vertebrae (2nd 15 mm), dorsal ribs, eleven rows of gastralia, thirteen caudal vertebrae (17 mm), coracoid, humerus (42 mm), manus, ilial fragment, pubis (95 mm), ischium (50 mm), femur (86 mm), tibia (104 mm), astragalus, metatarsal (65 mm), three partial metatarsals, pedal digit (20 mm), gastrolith(?)
Pliensbachian-Toarcian, Early Jurassic
Portland Formation, Connecticut, US
Referred- ?(BSNH 13656) (2.5-3 m) dorsal rib, dorsal rib fragments, pubis (248 mm), tibia (~230-270 mm) (Colbert and Baird, 1958)
Diagnosis- (after Colbert and Baird, 1958) dorsal neural spines anteroposteriorly shorter than Coelophysis bauri.
Comments- Colbert and Baird (1958) found two differences from Coelophysis bauri- dorsal neural spines anteroposteriorly shorter; ischium differently shaped. The first is a good distinction (assuming it's real), but the second certainly needs to be more precise. Podokesaurus may fall within the genus Coelophysis or be indeterminate at the level of Coelophysidae, but this is not formalized here pending redescription of C. bauri. It is a coelophysid based on the elongate dorsal centra.
References- Talbot, 1911. Podokesaurus holyokensis, a new dinosaur from the Triassic of the Connecticut Valley. Amer. Jour. Sci. 4 469-479, pl. iv, 6 text-figs.
Colbert and Baird, 1958. Coelurosaur bone casts from the Connecticut Valley Triassic. Am. Mus. Novitates 1901:1-11.

Pterospondylus Jaekel, 1913
P. trielbae Jaekel, 1913
Norian, Late Triassic
Trossingen Formation, Germany
Holotype- second dorsal vertebra (32 mm)
Comments- This species has been associated and synonymized with Procompsognathus in the past based only on the elongate centrum. The transverse processes are triangular, resembling Megapnosaurus more than Liliensternus and Dilophosaurus. Thus, Rauhut and Hungerbuhler (2000) suggest the taxon may be a coelophysid.
References- Jaekel, 1913.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.

"Syntarsus" "mexicanum" Hernandez, 2002
Toarcian, Early Jurassic
La Boca Formation, Mexico
Material- (IGM 6624) partial twelfth dorsal vertebra (13.1 mm), partial thirteenth dorsal vertebra (20 mm), partial synsacrum (61.5 mm- 16.2, 12.7, 9.6, 9.6, 13.4 mm), incomplete fused pelvis
?...(IGM 6625) posterior braincase, incomplete laterosphenoid, fragments
Diagnosis- (after Munter, 1999) ilia contact dorsally on midline (may be taphonomic).
Comments- Discovered in 1994, Munter (1999) found IGM 6624 to be sister to Megapnosaurus (as Syntarsus) based on the antitrochanter projecting into the acetabulum (though this is also present in Coelophysis and Liliensternus). It differs in having ilia contact sacral neural spines two and three. The skull fragments may not belong to the same individual or taxon and were thought to be a neoceratosair or maniraptoran by Munter due to their pneumatized paroccipital roots. However, this has since been found to exist in coelophysids such as "M." kayentakatae, making the possibility it belongs to the same individual more likely. Hernandez (2002) erected a new species Syntarsus mexicanum from the locality, which is probably based on IGM 6624 since no other coelophysoid specimens have been reported from there and Munter emphasized similarity to what he called Syntarsus. However, it is a nomen nudum for not including a diagnosis (ICZN 13.1.1) or a type specimen (ICZN 16.4.1). Hernandez also mentioned cranial elements of Ceratosaurus, which are probably IGM 6625 based on Munter's comparison to neoceratosaurs. Munter and Clark later (2006) described the specimens and included IGM 6624 in Carrano et al.'s (2002) matrix where it emerged sister to Coelophysis+Megapnosaurus (excluded due to lacking fused sacral neural spines), and Rauhut's (2003) matrix where it was in the clade of Coelophysis+Megapnosaurus (the latter a chimaera of rhodesiensis and kayentakatae). They considered IGM 6625 provisionally theropod. The only potential apomorphy noted in the material has been the ilia which meet at the midline, but this may be taphonomic. Munter and Clark note that the dorsally oriented acetabulum suggests distortion, but the rugose medial ilial surface suggesting contact with the other ilium and general lack of distortion in other fossils from the locality suggests it was natural. As "mexicanum" has not been shown to be closer to Megapnosaurus than to Coelophysis, it is here retained under the name "Syntarsus" "mexicanum".
References- Clark, Montellano, Hopson, Hernandez and Reynoso, 1998. The Jurassic vertebrates of Huizachal canyon, Tamaulipas. Avances en Investigacion, Paleontologia de Vertebrados. Universidad Autonoma del Estado de Hidalgo Publicacion Especial 1. 1-3.
Munter, 1999. Two theropod dinosaur specimens from Huizachal Canyon, Mexico. Journal of Vertebrate Paleontology. 19(3), 65A.
Munter, 1999. Two theropod dinosaur specimens from Huizachal Canyon, Mexico. Masters thesis. George Washington University.
Hernandez, 2002. Los dinosaurios en Mexico. in Gonzalez Gonzalez and De Stefano Farias (eds.). Fosiles de Mexico: Coahuila, una Ventana a Traves del Tiempo. Gobierno del Estado de Coahuila, Saltillo. 143-153.
Munter and Clark, 2006. Theropod dinosaurs from the Early Jurassic of Huizachal Canyon, Mexico. In Carrano, Gaudin, Blob, Wible (eds.). Amniote paleobiology: Perspectives on the evolution of mammals, birds, and reptiles. University of Chicago Press, Chicago. 53-75.

Segisaurinae Camp, 1936 sensu Kalandadze and Rautian, 1991
Definition- (Segisaurus halli <- Coelophysis bauri) (proposed)
Diagnosis- (after Tykoski, 2005) anterodorsal margin of acromion has smooth, continuous, high-angle transition to scapular blade; pubes rectangular in distal view.
Comments- This subfamily was based on Camp's family Segisauridae and is used here for the clade formed by Segisaurus, "Megapnosaurus" kayentakatae and possibly Camposaurus recovered by Tykoski (2005). Such a clade was not recovered by Ezcurra and Novas (2006), where "Megapnosaurus" kayentakatae was instead closer to Coelophysis and Megapnosaurus than to Segisaurus.
References- Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of Arizona. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 24, 39-56.
Kalandadze and Rautian, 1991. Late Triassic zoogeography and reconstruction of the terrestrial tetrapod fauna of North Africa. Paleont. Journ. 1, 1-12.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.

Camposaurus Hunt, Lucas, Heckert, Sullivan and Lockley, 1998
C. arizonensis Hunt, Lucas, Heckert, Sullivan and Lockley, 1998
Early Norian, Late Triassic
Mesa Redondo Member of the Chinle Formation, Arizona, US
Holotype- (UCMP 34498) distal tibiae, distal fibulae, astragalocalcanea
Referred- ?(UCMP 25791) pubic fragment, ilial fragment (Hunt, Lucas, Heckert, Sullivan and Lockley, 1998)
Diagnosis- indeterminate relative to "Megapnosaurus" kayentakatae.
Comments- Originally referred to as Ceratosauria indet. by Long and Murray (1995), UCMP 34498 was made the holotype of a new coelophysid taxon by Hunt et al. (1998). Downs (2000) suggests that Camposaurus is a junior synonym of Coelophysis, since it falls within the range of variation of the Ghost Ranch specimens, and the initial comparisons were made with Colbert's inaccurate monograph. Nesbitt et al. (2005, 2007) agree it cannot be distinguished from Coelophysis bauri, as the straight ventral astragalar margin is found in the latter as well.. Tykoski (2005) notes the presence of fibulocalcanear fusion is otherwise only seen in "Megapnosaurus" kayentakatae, though otherwise the material cannot be placed more precisely within the Coelophysis<Liliensternus clade.
UCMP 25791 was figured by Long and Murray, then assigned questionably to Camposaurus by Hunt et al.. Tykoski noted the pelvic fragment indicates the ilium was fused to the pubis, which is a ceratosaurian character. Thus it may be properly referred to Camposaurus.
A proximal femur (UCMP 139662), dorsal centra (MNA V3091; UCMP 177317), and sacral remains (UCMP 138591, 178047-178049) were identified as Ceratosauria indet. by Long and Murray, then made paratypes of Camposaurus by Hunt et al.. The latter authors felt the material may even belong to the holotype individual, but Nesbitt et al. ciuld only identify the femur to Saurischia indet. and the axial material to Archosauria indet..
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the western United States. Geobios, 31(4):511-531.
Downs, 2000. Coelophysis bauri and Syntarsus rhodesiensis compared, with comments on the perparation and preservation of fossils from the Ghost Ranch Coelophysis quarry. in Lucas and Heckert (eds.), 2000. Dinosaurs of New Mexico. NMMNH Bulletin 17. 33-37.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. JVP. 25(3) 96A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

"Megapnosaurus" kayentakatae (Rowe, 1989) Tykoski and Rowe, 2004
= Syntarsus kayentakatae Rowe, 1989
= Coelophysis kayentakatae (Rowe, 1989) Bristowe and Rowe, 2004
Hettangian, Early Jurassic
Silty Facies Member of the Kayenta Formation, Arizona, US
Holotype- (MNA V2623) (robust adult) skull (230 mm), mandibles, hyoids, atlas, axis (27.8 mm), third cervical vertebra (33 mm), fourth cervical vertebra (>41 mm), fifth cervical vertebra, sixth cervical vertebra (>45 mm), seventh cervical vertebra (46.7 mm), eighth cervical vertebra (43.5 mm), ninth cervical vertebra, tenth cervical vertebra (>33 mm), cervical ribs 1-10 (110-295 mm), first dorsal vertebra, partial second dorsal vertebra, first dorsal rib, over fifteen rows of gastralia, two partial sacral centra, caudal vertebrae, scapulocoracoids (one partial; 181 mm), furcula, incomplete humerus (~116 mm), distal carpal I+II, proximal carpal, carpal, metacarpal I (21.2 mm), manual ungual I (21 mm), phalanx II-2 (24 mm), proximal manual ungual II, metacarpal III (>39 mm), phalanx III-3 (18.8 mm), manual ungual III (18.2 mm), metacarpal IV (>20.8 mm), phalanx IV-1 (13 mm), phalanx IV-2, partial pubes, partial ischia, femora (one partial) (276 mm), tibiae (one partial) (292 mm), fibulae (one partial) (274 mm), astragalocalcaneum (32.2, 33.5 mm transversely), distal tarsal III, distal tarsal IV, metatarsal I (38 mm), phalanx I-1 (27.2, 27.3 mm), pedal ungual I (15, 16 mm), metatarsal II (150 mm), phalanx II-1 (47, 44.8 mm), phalanx II-2 (38 mm), pedal ungual II (30 mm), metatarsal III (173 mm), phalanx III-1 (50, 47.7 mm), phalanx III-2 (33, 36.9 mm), phalanx III-3 (34, 34.6 mm), pedal ungual III (28.5 mm), metatarsal IV, phalanx IV-1 (27.5 mm), phalanx IV-2, phalanx IV-3 (18.3, 18.4 mm), phalanx IV-4 (15.5, 15.6 mm), pedal ungual IV (>24 mm), metatarsal V (>57 mm)
Paratypes- (MNA V2623) (15-20% larger than holotype; at least two robust) snout, dentaries, maxillary fragment with ninth to eleventh teeth, frontals, partial occipital condyle, fragments of sacra, portions of ~34 caudal centra, two fragmentary femora, three proximal tibiae, three astragalocalcanea, eight ends of metatarsals
(MCZ 8817) (at least eleven robust individuals) caudal, pelvic and hindlimb elements
?(MNA V100) partial ilium
?(MNA V140) (juvenile) humerus, femur, partial tibia, bone fragments
(TMM 43648-9)
(TMM 43669-3) (young subadult)
(TMM 43688-1) (robust old subadult) cervical vertebrae, sacrum (27, 23.4, 22.2, 24, 26 mm = 121 mm), few caudal centra, partial ilia, pubes (<232 mm), ischia (121 mm), femora (229.8, 230 mm), proximal tibia, proximal fibula, distal tarsal III, distal tarsal IV, proximal tarsometatarsus, several pedal phalanges
Diagnosis- (after Tykoski and Rowe, 2004) nasal crests; frontals separated by midline anterior extension of parietals.
Other diagnoses- Tykoski and Rowe (2004) listed transverse groove on anterior astragalar surface as an additional diagnostic feature of this species, but it is also present in Coelophysis bauri and the Shake-n-Bake coelophysid (Tykoski, 2005).
Comments- The 'juveniles' previously referred (Rowe, 1989) are actually the distinct "Shake-n-Bake" coelophysid (Tykoski, 1998). The sacral and caudal material assigned to the holotype may belong to the other two individuals with the same specimen number. Tykoski (2005) found this species to be more closely related to Segisaurus than to Coelophysis or Megapnosaurus.
UCMP V128659 was discovered in 1982 and referred to Syntarsus kayentakatae by Rowe (1989), as a subadult gracile individual. Tykoski (1998) did not examine it for his redescription of the species, but later (2005) examined it for his PhD thesis and considered it to be "probably referrable to "Syntarsus" kayentakatae" without discussion. Gay (2010) described the specimen as the new tetanurine taxon Kayentavenator elysiae. However, the description and analysis contain numerous errors, and a placement in Coelophysidae seems correct, though there is no published evidence Kayentavenator is the same taxon as kayentakatae.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9, 125-136.
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications for ceratosaurid phylogeny. Unpublished Masters Thesis, University of Texas at Austin, 217 pp.
Ivie, Slipinski and Wegrzynowicz, 2001. Generic homonyms in the Colydiinae (Coleoptera: Zopheridae). Insecta Mundi 15:63-64.
Tykoski, Forster, Rowe, Sampson and Munyikwa, 2002. A furcula in the coelophysid theropod Syntarsus. Journal of Vertebrate Paleontology. 22(3):728–733.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Gay, 2010. Notes on Early Mesozoic theropods. Lulu Press. 44 pp.

Segisaurus Camp, 1936
S. halli Camp, 1936
Toarcian, Early Jurassic
Navajo Sandstone, Arizona, US
Holotype- (UCMP 32101) (robust old subadult) (1.54 m) distal cervical ribs 7-10, partial fourth dorsal vertebra, partial fifth dorsal vertebra, partial sixth dorsal vertebra, partial eighth dorsal vertebra, partial ninth dorsal vertebra, dorsal neural arch, several partial dorsal ribs, over twelve rows of gastralia, impression of first sacral centrum, impression of second sacral centrum, partial third sacral vertebra, partial fouth sacral vertebra, partial caudal vertebrae 2-22, fragments of several caudal vertebrae, several chevrons, scapula (93 mm), partial coracoids, partial furcula, possible sternal fragment, incomplete humerus (72 mm), proximal radius, proximal ulna, distal phalanx II-2, manual ungual II, distal phalanx III-2, phalanx III-3, manual ungual III, manual phalanges, ilial fragments, incomplete pubes (~110 mm), incomplete ischia (~96 mm), incomplete femora (~145 mm), tibiae (160 mm), incomplete fibula, fibular fragment, partial astragalus, calcaneum, distal tarsal IV, metatarsal I (33 mm), phalanx I-1 (17 mm), pedal ungual I, metatarsals II, phalanx II-1, phalanx II-2 (23 mm), pedal ungual II, metatarsals III (99 mm), phalanx III-1, phalanx III-2 (22 mm), phalanx III-3 (13 mm), pedal ungual III (10 mm), metatarsal IV, phalanx IV-1, phalanx IV-2 (14 mm), phalanx IV-3, phalanx IV-4, pedal ungual IV (14 mm), metatarsals V (32 mm)
Diagnosis- (from Rauhut, 2003) dorsal centra not very constricted ventrally; slender scapula; humeral shaft with stronger torsion than Coelophysis (~50 degrees); large ischial fenestra; (from Carrano et al., 2005) rectangular humeral deltopectoral crest.
Comments- Segisaurus has been reprepared, showing a furcula in articulation with the coracoids (not separate clavicles) and thin-walled long bones.
Although Carrano et al. (2005) could not recover any resolution within Coelophysoidea in their cladistic analysis, when several characters are changed to ordered, the results differ. If a standard coelophysoid topology of (Dilophosaurus (Liliensternus (Coelophysis, Megapnosaurus))) is enforced, Segisaurus is placed outside the Liliensternus + Coelophysis + Megapnosaurus clade. This differs from Rauhut's (2000) findings, which placed it in the Coelophysidae.
References- Camp and VanderHoof, 1935. Small bipedal dinosaur from the Jurassic of northern Arizona. Proc. Geol. Soc. Amer. 1934 384-385.
Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of Arizona. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 24, 39-56.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Senter and Hutchinson, 2001. New information on the skeleton of the theropod Segisaurus halli. JVP 21(3) 100A.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate Paleontology. 25(4), 835–849.

Coelophysinae Nopcsa, 1928
Definition- (Coelophysis bauri <- Segisaurus halli) (proposed)
Other definitions- (Coelophysis bauri <- Procompsognathus triassicus) (Sereno, in press; modified from Sereno, 1998)
= Syntarsiinae Kalandadze and Rautian, 1991
Diagnosis- (modified from Tykoski, 2005) dorsal process of maxilla angles <35 degrees from horizontal; promaxillary fenestra absent; lateral lamina of ventral lacrimal process linear and remains posterior to medial lamina; more than 18 dentary teeth; jugal does not contact antorbital fossa.
Comments- Tykoski (2005) also included the reduction in size of dorsolateral nasal ridges as a synapomorphy, but as Zupaysaurus' ridges are more likely to be displaced nasals, this becomes an ambiguous character not favored by DELTRAN. It is viewed as evolving convergently in Dilophosaurus and "Megapnosaurus" kayentakatae here.
Alhough originally proposed by Nopcsa in 1928 to only include Coelophysis (and not Podokesaurus, Procerosaurus, Saltopus or Tanystropheus), this subfamily went largely unused until Paul (1988) used it for Coelophysis, Megapnosaurus rhodesiensis and Elaphrosaurus, to separate them from the halticosaurine Liliensternus and Dilophosaurus. Sereno (1998) then used it to group Coelophysis and Megapnosaurus together, defining it to include Coelophysis and exclude Procompsognathus. In Sereno's topology, this also excluded Segisaurus and Liliensternus. Yet subsequent studies (e.g. Tykoski, 2005; Ezcurra and Rautian, 2006) have shown Procompsognathus to have an uncertain placement within derived coelophysoids, making the precise application of Sereno's definition impossible and the content of his Coelophysinae (besides Coelophysis itself) unknown. Carrano et al. (2002) later used Coelophysinae for a Megapnosaurus + Coelophysis clade, excluding Liliensternus. Taking these prior uses into account, Coelophysinae is here used for the Megapnosaurus + Coelophysis clade, but with Segisaurus as the external specifier instead of Procompsognathus. This is a clade recovered in most phylogenetic analyses where coelophysid interrelationships are resolved (e.g. Tykoski, 2005; Ezcurra and Novas, 2006) and is equivalent to Sereno's Coelophysinae in content.
Kalandadze and Rautian (1991) proposed the subfamily Syntarsiinae, but this cannot be used for a theropod group, as Syntarsus is the name of a beetle (Ivie et al., 2001).
References- Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Kalandadze and Rautian, 1991. Late Triassic zoogeography and reconstruction of the terrestrial tetrapod fauna of North Africa. Paleont. Journ. 1, 1-12.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 210(1), 41-83.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology. 22(3), 510-534.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article, 38 pp. DOI 10.1080/08912960600845791.

Coelophysidae sensu Tykoski and Rowe, 2004
Definition- (Coelophysis bauri + Megapnosaurus rhodesiensis) (modified)

Coelophysinae sensu Sereno, 1998
Definition- (Coelophysis bauri <- Procompsognathus triassicus)
Coelophysis Cope, 1889
= Rioarribasaurus Hunt and Lucas, 1991
C. bauri (Cope, 1887a) Cope, 1889
= Coelurus bauri Cope, 1887a
= Coelurus longicollis Cope, 1887a
= Tanystropheus bauri (Cope, 1887a) Cope, 1887b
= Tanystropheus longicollis (Cope, 1887a) Cope, 1887b
= Coelophysis longicollis (Cope, 1887a) Cope, 1889
= Rioarribasaurus colberti Hunt and Lucas, 1991
= Syntarsus colberti (Hunt and Lucas, 1991) Paul, 1993
Late Rhaetian-Early Hettangian, Late Triassic-Early Jurassic
Siltstone member of the Chinle Formation, New Mexico, US
Neotype - (AMNH 7224; holotype of Rioarribasaurus colberti) (2.86 m, 19.9 kg) complete skeleton including skull (191 mm), axis (31 mm), cervical series (405 mm), dorsal series (455 mm), scapula (131 mm), humerus (134 mm), radius (81 mm), ulna (72 mm), manual ungual I (21 mm), metacarpal III (40 mm), pubis (233 mm), femur (203 mm), tibia (221 mm), astragalus, calcaneum, (astragalocalcaneum 23 mm wide), metatarsus (120 mm) (Colbert and Baird, 1958)
Referred- (AMNH 7223) (2.68 m, 15.3 kg) complete skeleton including skull (265 mm), cervical series (485 mm), dorsal series (425 mm), humerus (120 mm), femur (209 mm), tibia (224 mm), astragalus, calcaneum, metatarsal II (110 mm), metatarsal III (126 mm), metatarsal IV (114 mm) (Colbert and Baird, 1958)
(AMNH 7225) skull (Colbert, 1989)
(AMNH 7226) pes (Colbert, 1989)
(AMNH 7227) skull (159 mm), cervical series (384 mm), dorsal series (360 mm), sacrum, scapulae, humeri (86 mm), radii, ulnae, manus, ilium, pubis (145.4 mm), proximal ischium, femora (165 mm), tibiae (151.8 mm), fibulae, pes (Colbert and Baird, 1958)
(AMNH 7228) skull (198 mm), cervical series (430 mm), dorsal series (330 mm), sacrum, scapula, humerus (89 mm), radius, ulna, ilium, pubes, ischium, femora (164 mm), tibiae (188 mm), fibulae, metatarsus (108 mm), pedal phalanges (Colbert, 1964)
(AMNH 7229) six dorsal vertebrae, sacral vertebrae, caudal vertebrae, humerus (75 mm), partial ilia, pubis (125 mm), ischium, femora (135 mm), tibiae (154 mm), fibulae, metatarsus (85 mm), pedal phalanges (Colbert and Baird, 1958)
(AMNH 7230) skull (110.5 mm), cervical series (215 mm), dorsal series (225 mm), sacrum, sacpulae, coracoids, humeri (61 mm), radius, ulna, manus, ilium, pubis (113 mm), femora (125 mm), tibiae (123.9 mm), fibulae (Colbert and Baird, 1958)
(AMNH 7231) incomplete cervical series (215 mm), incomplete dorsal series (262 mm), sacrum, scapulae, coracoids, humerus (58 mm), radius, ulna, partial ilium, pubis (139.8 mm), femora (132 mm), tibiae (156 mm), fibulae, pes (Colbert and Baird, 1958)
(AMNH 7232) humerus (57 mm), ilium, pubes, femur (141 mm), tibia (157 mm), fibula, metatarsus (95 mm), pedal phalanges (Colbert and Baird, 1958)
(AMNH 7233) dorsal vertebrae, sacral vertebrae, ilium, pubis (125 mm), ischium, femora (126 mm), tibiae (140 mm), fibulae, metatarsus (81 mm), pedal phalanges (Colbert and Baird, 1958)
(AMNH 7234) dorsal vertebrae, sacral vertebrae, partial pubis, ischium, femur (118 mm), tibia (135 mm), fibula, metatarsus (84 mm), pedal phalanges (Colbert, 1964)
(AMNH 7235) ilium, ischium, partial femur (Colbert, 1989)
(AMNH 7236) ilium, pubis (122 mm), femur, tibia (134.1 mm) (Colbert and Baird, 1958)
(AMNH 7237) skull, cervical vertebrae (Colbert, 1989)
(AMNH 7238) skull, humerus (55 mm), femur (126 mm), tibia (147 mm), metatarsus (87 mm) (Colbert, 1964)
(AMNH 7239) skull, mandibles, cervical vertebrae (Colbert, 1989)
(AMNH 7240) skull (198 mm), mandibles (Colbert, 1989)
(AMNH 7241) skull (140 mm), mandibles (Colbert, 1989)
(AMNH 7242) (juvenile) skull (~68 mm) (Colbert, 1989)
(AMNH 7243) ten dorsal vertebrae, dorsal ribs, sacrum, proximal caudal vertebrae, scapulacoracoid, humeri (91 mm), radii, ulnae, manus, pelvis, femora (172 mm), tibiae (199 mm), fibulae, astragalus, calcaneum, metatarsus (114 mm), pedal phalanges (Colbert, 1964)
(AMNH 7244) proximal thirty caudal vertebrae, ilium, pubis (170 mm), ischium, femora (173 mm), tibiae (195 mm), fibulae, metatarsus (120 mm), pedal phalanges (Colbert and Baird, 1958)
(AMNH 7245) five or six presacral vertebrae, six proximal caudal vertebrae, ilium, pubis (230 mm), ischium, femur, tibia (228 mm) (Colbert and Baird, 1958)
(AMNH 7246) eight caudal vertebrae, pelvis, femur (122 mm), tibiae (136 mm), fibulae, metatarsi (79 mm), pedal phalanges (Colbert, 1964)
(AMNH 7247) femur (125 mm), tibia (138 mm), metatarsus (84 mm), pedal phalanges (Colbert, 1964)
(AMNH 7248) ilium, pubis (Colbert, 1989)
(AMNH 7249) eight dorsal vertebrae, dorsal ribs, sacrum, five caudal vertebrae, ilium, pubis (220 mm), ischium, femora (196 mm), tibiae (207 mm), fibulae, metatarsi (110 mm), pedal phalanges (Colbert and Baird, 1958)
(AMNH 7250) distal tibia, distal fibula, pes (Colbert, 1989)
(AMNH 7251) three dorsal vertebrae, sacrum, six or more caudal vertebrae, ilium, pubes, femur, tibiae, fibula, pes (Colbert, 1989)
(AMNH 7252) seven presacral vertebrae, sacrum, two or more caudal vertebrae, pelvis, hindlimb (Colbert, 1989)
(AMNH 7253) tibia (155 mm), fibula, metatarsus (91 mm), pedal phalanges (Colbert, 1964)
(AMNH 7254) eleven caudal vertebrae (Colbert, 1989)
(AMNH 7255) partial skull (Colbert, 1989)
(AMNH 7256) tibia (152 mm), fibula, metatarsi (82 mm), pedal phalanges (Colbert, 1964)
(AMNH 7257) seventeen presacral vertebrae, humeri, partial radii, partial ulnae (Colbert, 1989)
(AMNH 7258) skull, mandible, cervical vertebrae 1-7, seven or eight dorsal vertebrae (Colbert, 1989)
(CM 31374) skull, mandible (Downs, 2000)
(CM C-1-82) (one or two individuals) partial skeleton (Colbert, 1989)
(CM C-3-82) (juvenile) skeleton including nasal, vertebrae, furcula, pelvis, hindlimbs (Downs, 2000)
(CM C-4-81) (at least nine individuals) skull (250 mm), maxillary fragment, caudal vertebrae 1-5, two pelves, two hindlimbs, femur, distal tibiae, two pes (Colbert, 1989)
(GR141) (Downs, 2000)
(GR142) (Downs, 2000)
(GR1442) (Downs, 2000)
(MCZ 4326) skull, mandible (Colbert, 1989)
(MCZ 4327) skull (239 mm), mandible, atlas, axis, pubis, femur, proximal tibia (Colbert, 1989)
(MCZ 4328) partial maxilla (Colbert, 1989)
(MCZ 4329) forelimb including five carpals (Colbert, 1989)
(MCZ 4330) pelvis (Colbert, 1989)
(MCZ 4331) (first individual) last thirteen dorsal vertebrae, dorsal ribs, humerus, radius, ulna, pelvis, hindlimbs
(second individual) two distal caudal vertebrae, femur, tibia, other elements
(third individual) three cervical vertebrae, three dorsal vertebrae, pelvis, femora, tibia (Colbert, 1989)
(MCZ 4332) (three or four individuals) vertebrae, pelves, hindlimbs (Colbert, 1989)
(MCZ 4333) skull, partial mandible, anterior cervical vertebrae, partial manus (Colbert, 1989)
(MCZ 4334) vertebrae, pelvis, hindlimbs (Colbert, 1989)
(MCZ 4335) vertebrae (Colbert, 1989)
(MNA V3139) fragmentary skull, fragmentary mandibles, postcrania (Colbert, 1989)
(MNA V3315) skull (143 mm), mandibles, atlas, axis, third cervical vertebra (Colbert, 1989)
(MNA V3318) (1.8 m) incomplete skeleton including skull (~88 mm), cervical series (184 mm), dorsal series (255 mm), femur (123 mm), tibia (136 mm), astragalocalcaneum, metatarsal II (72 mm), metatarsal III (82 mm), metatarsal IV (70.5 mm) (Colbert, 1989)
(MNA V3319) pes including metatarsal II (67 mm), metatarsal III (76 mm), metatarsal IV (69 mm) (Colbert, 1989)
(MNA V3320) pes including astragalus, calcaneum, metatarsal II (115 mm), metatarsal III (128 mm), metatarsal IV (118 mm) (Colbert, 1989)
(MNA V3321) distal tibia, astragalocalcaneum (Colbert, 1989)
(MNA V3322) partial skull (Colbert, 1989)
(MNA V3323) caudal series (Colbert, 1989)
(NMMNH P-42352) skeleton including cololite and coprolite (Rinehart, Hunt, Lucas, Heckert and Smith, 2005)
(NMMNH P-44552) skeleton including coprolite with juvenile elements included (rib fragments, ulnare, proximal metacarpals, partial phalanges, long bone fragments, cranial and/or pelvic material) (Rinehart, Hunt, Lucas, Heckert and Smith, 2005)
(NMMNH P-44801) skeleton including coprolite (Rinehart, Hunt, Lucas, Heckert and Smith, 2005)
(NMMNH P-C-8-82) (several individuals including three juveniles) material including five furculae (Rinehart, Lucas and Hunt, 2006)
(RTMP 84-63-23) skull (Colbert, 1989)
(RTMP 84-63-29) skull, mandibles, cervical vertebrae, dorsal vertebrae (Colbert, 1989)
(RTMP 84-63-30) cervical vertebrae, dorsal vertebrae, scapula, coracoid, humerus, radius, ulna, manus (Colbert, 1989)
(RTMP 84-63-31) maxilla, mandible (Colbert, 1989)
(RTMP 84-63-32) anterior skull, anterior mandible, posterior cervical vertebrae, dorsal vertebrae, ribs, scapulocoracoid, forelimbs (Colbert, 1989)
(RTMP 84-63-33) last thirteen dorsal vertebrae, ribs, sacrum, more than fifteen caudal vertebrae, scapula, coracoid, humerus, pelvis, hindlimbs including metatarsal II (94.2 mm), metatarsal III (105 mm), metatarsal IV (93 mm) (Colbert, 1989)
(RTMP 84-63-34) last seven dorsal vertebrae, several caudal vertebrae, pelvis, femora, tibiae, fibula, astragalocalcaneum, pes (Colbert, 1989)
(RTMP 84-63-35) eleven caudal vertebrae, humerus(?), tibia, fibula, astragalus, calcaneum, pes (Colbert, 1989)
(RTMP 84-63-36) vertebrae, femora, tibiae, other elements (Colbert, 1989)
(RTMP 84-63-37) vertebrae, scapulacoracoid(?), ischia, hindlimbs (Colbert, 1989)
(RTMP 84-63-38) pelvis (Colbert, 1989)
(RTMP 84-63-39) vertebrae, ilium, femur (Colbert, 1989)
(RTMP 84-63-40) manus (Colbert, 1989)
(RTMP 84-63-41) six caudal vertebrae (Colbert, 1989)
(RTMP 84-63-42) partial femora, tibiae (Colbert, 1989)
(RTMP 84-63-43) seven series of caudal vertebrae (Colbert, 1989)
(RTMP 84-63-44) vertebrae, other elements (Colbert, 1989)
(RTMP 84-63-45) fragmentary skull (Colbert, 1989)
(RTMP 84-63-46) manus (Colbert, 1989)
(RTMP 84-63-47) distal tibia, distal fibula, astragalocalcaneum (Colbert, 1989)
(RTMP 84-63-48) sacrum, posterior ilia (Colbert, 1989)
(RTMP 84-63-49) eleven vertebrae (Colbert, 1989)
(RTMP 84-63-50) manus (Colbert, 1989)
(RTMP 84-63-51) dorsal vertebrae, dorsal ribs (Colbert, 1989)
(RTMP 84-63-52) carpus, manus (Colbert, 1989)
(RTMP 84-63-53) seven cervical vertebrae (Colbert, 1989)
(RTMP 84-63-54) ten caudal vertebrae (Colbert, 1989)
(RTMP 84-63-55) femur (Colbert, 1989)
(RTMP 84-63-56) fourth distal tarsal, metatarsal (Colbert, 1989)
(RTMP 84-63-57) vertebrae (Colbert, 1989)
(RTMP 84-63-58) caudal vertebrae, long bone (Colbert, 1989)
(RTMP 84-63-59) six posterior cervical vertebrae (Colbert, 1989)
(RTMP 84-63-60) element (Colbert, 1989)
(RTMP 84-63-61) vertebrae, two metatarsals (Colbert, 1989)
(YPM 41196) skull (211 mm), mandible (Colbert, 1989)
(YPM 41197) pelvis (Colbert, 1989)
(YPM 41412) pelvis, hindlimb (Colbert, 1989)
(YPM 43506) anterior skull (Colbert, 1989)
Late Norian, Late Triassic
Petrified Forest Member of the Chinle Formation, Arizona, New Mexico, US
Material- (AMNH 2701; lectotype of Coelurus longicollis) cervical vertebra (63 mm) (Cope, 1887a)
?(NMMNH P-29046) femora (155.5 mm), tibia (164 mm), proximal tibia, proximal fibula (Heckert et al., 2000)
?(NMMNH P-29047) manual ungual, metacarpal IV, partial ilium, incomplete ischium (115.3 mm), proximal tibia, proximal fibula, distal metatarsal, proximal pedal phalanx, phalanx IV-? (Heckert et al., 2000)
?(NMMNH P-30779) dorsal vertebra (Heckert et al., 2000)
?(NMMNH P-30780) dorsal vertebra (Heckert et al., 2000)
(NMMNH P-30852) premaxilla, maxilla, lacrimal, prefrontal, postorbital, incomplete mandibles (133.9, 122.4 mm), hyoids, third cervical vertebra, fourth cervical vertebra, cervical ribs (Heckert et al., 2000)
?(NMMNH P-31293) cervical rib, incomplete tibia (Heckert et al., 2000)
?(NMMNH P-31661) cervical ribs, incomplete sacrum, partial scapulocoracoid, radius (51.8 mm), metacarpal, pedal ungual (Heckert et al., 2000)
?(NMMNH P-54617) tibia (Spielmann et al., 2007)
?(NMMNH P-54618) distal femur (Spielmann et al., 2007)
?(NMMNH P-54619) distal femur (Spielmann et al., 2007)
?(NMMNH P-54620) femur (245 mm) (Spielmann et al., 2007)
(PEFO 33981) fragmentary skeleton including partial posterior vertebral column, pelvis, incomplete hindlimbs (Nesbitt et al., 2007)
(PEFO 33983) incomplete posterior skeleton (Nesbitt et al., 2007)
(UCMP 129618) (3.46 m) dorsal centrum (45 mm), partial dorsal centrum, three partial sacral centra, proximal caudal centrum (33 mm), partial ilium, nearly complete pubis, partial ischium, femora (235, 245 mm), tibia (255 mm), fibula, astragalocalcaneum, distal tarsal III, distal tarsal IV, phalanx I-1 (29 mm), metatarsal III (150 mm), proximal phalanx III-1, phalanx III-2 (37 mm), distal pedal ungual III, metatarsal IV (133 mm), phalanx IV-1 (30 mm), phalanx IV-3 (22 mm), phalanx IV-4 (19 mm), proximal pedal ungual IV (Padian, 1986)
partial skeleton including teeth, cervical vertebrae, femora and tibiae (Hunt and Wright, 1999)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, Texas, US
?(TTUP 10071) partial ilium (Lehman and Chatterjee, 2005)
?(TTUP 10072) vertebrae, proximal femur, distal tibia, astragalus (Lehman and Chatterjee, 2005)
?(TTUP 10082) pubis (Lehman and Chatterjee, 2005)
Late Triassic(?)
US(?)
(CM 11894) tibia, fibula, astragalus, calcaneum (Tykoski, 2005)
(SMP VP-453) (Sullivan et al., 1996)
(SMP VP-456) (Sullivan et al., 1996)
(SMP VP-462) (Sullivan et al., 1996)
(SMP VP-465) (Sullivan et al., 1996)
(SMP VP-469) (three individuals) (Sullivan et al., 1996)
(SMP VP-476) (Sullivan et al., 1996)
(SMP VP-478) (four specimens) (Sullivan et al., 1996)
(SMP VP-487) (Sullivan et al., 1996)
(SMP VP-491) (Sullivan et al., 1996)
(SMP VP-1072) femur
(TMM 45559) cervical vertebrae, sacral vertebrae, partial ilium, femur, tibia, astragalocalcaneum (Tykoski, 2005)
Removed material- Cope (1887a) referred to a Coelophysis tooth, and Padian (1986) noted that teeth preserved in his material (specifically AMNH 2733) belong to parasuchians. The distal femur AMNH 2725 belongs to a shuvosaurid (Nesbitt et al., 2007). The distal femur AMNH 2721 was originally referred to bauri, then to longicollis, but is Dromomeron romeri (Nesbitt et al., 2009). Colbert and Baird (1958) referred BSNH 13656 from the Portland Formation of Connecticut to Coelophysis sp., but these were later referred to holyokensis by Colbert (1964). Several authors (Gregory, 1945; Elder, 1978, 1987) assigned material from the Colorado City Member of the Dockum Formation in Texas to Coelophysis, but these are Trilophosaurus (Hunt et al., 1998). The vertebral centra from the Blue Mesa Member of the Chinle Formation referred to Coelophysis by Colbert (1989) are Trilophosaurus (Hunt et al., 1998).
Diagnosis- (after Tykoski, 1998) anterior pedunclar foramina in cervical vertebrae (unknown in Megapnosaurus).
(after Downs, 2000) unspecified differences from Megapnosaurus in cervical length, proximal and distal hindlimb proportions and proximal caudal vertebral anatomy.
(after Tykoski and Rowe, 2004) differs from Megapnosaurus and "M." kayentakatae in lacking pit at the base of the nasal process of the premaxilla.
(after Bristowe and Raath, 2004) differs from Megapnosaurus in having a longer maxillary tooth row; anteroposterior length of ventral lacrimal process >30% of its height.
(after Ezcurra, 2007) absence of an offset rostral process of the maxilla; strongly caudally bowed quadrate; small external mandibular fenestra (9-10% of mandibular length) (also in Dilophosaurus). Differs from Megapnosaurus in having a square-shaped rostral process of the jugal; unreduced medial condyle of the distal trochlea of the metacarpal I.
Other diagnoses- Cope (1887a) originally diagnosed Coelurus bauri based on several characters. Posterior pleurocoels in the cervical centra are present in all coelophysoids, including the longicollis type. The dorsal longitudinal grooves on the anterior cervical neural arch (alongside the neural spine) are present in other coelophysoids as well. The smaller size than longicollis is within the range of ontogenetic or individual variation. The femur is said to be "not so strongly grooved at the third [=fourth] trochanteric ridge", but this specimen (AMNH 2725) has been reassigned to a shuvosaurid by Nesbitt et al. (2007).
Cope (1889) erected Coelophysis because the vertebrae have neural canals, which he did not believe were present in Tanystropheus. Besides being primitive, this is untrue for Tanystropheus. In addition, Cope noted the amphicoelous cervicals differed from Coelurus (though the opisthocoelous cervicals assigned to Coelurus by Marsh have since been removed) and the lack of an ectocondylar tuber on the femur differed from Megadactylus (=Anchisaurus) (though this was based on a silesaurid femur).
Colbert (1964) listed numerous characters in his diagnosis of Coelophysis, most of which are primitive for avepods- lightly built; hollow bones; skull narrow; teeth laterally compressed and serrated; amphicoelous vertebrae; ten cervical vertebrae; thirteen dorsal vertebrae; five sacral vertebrae; sacral vertebrae fused; distal caudals elongate; forelimb ~50% of hindlimb length; carpals present; manual digit IV reduced; elongate ilium; pubis anteroposteriorly flattened; pubic boot; ischium rod-like; ischium proximally expanded. Some are not true in Coelophysis- pubis equal or longer than femur.
Comments- The original type material was discovered in 1881 and described by Cope (1887a) as Coelurus bauri and C. longicollis, though they were referred to that genus without justification. C. bauri's syntypes are a partial cervical vertebra (AMNH 2717), an incomplete sacrum and pubic peduncle of the ilium (AMNH 2722) and a distal femur now referred to Shuvosauridae by Nesbitt et al. (2007) (AMNH 2725). Cope later (1887b) referred these to Tanystropheus instead, along with a third species from the same collection- T. willistoni. He felt the amphicoelous cervicals of his material were more similar to the amphicoelous vertebrae of Tanystropheus (then thought to be caudals) than the supposedly opisthocoelous cervicals of Coelurus (based on vertebrae now removed from that taxon- YPM 1996 and 1997). He referred additional material (AMNH 2718-2721, 2723 and 2724) to C. bauri based on size. In 1889, Cope separated the three species from Tanystropheus as his new genus Coelophysis. Huene (1906, 1915) illustrated the material and described it in more depth. In 1906 Huene referred the caudal AMNH 2744 to C. bauri, while in 1915 he reassigned numerous specimens. The syntype dorsal of C. longicollis AMNH 2715 was reassigned to C. bauri, and the pubic peduncle in C. bauri's syntype AMNH 2722 was assigned to C. willistoni. Additionally AMNH 2708 was reassigned to C. bauri, while AMNH 2723 and 2744 were reassigned to C. willistoni, and AMNH 2721 was reassigned to longicollis. AMNH 2750, 2752 and perhaps 2745 and 2751 were also assigned to bauri. These referrals were all based on size, and there is no reason to believe they all belong to coelophysoids, let alone a single species.
longicollis- Coelurus longicollis was originally named by Cope (1887a) based on a cervical vertebra (AMNH 2701), dorsal vertebra (AMNH 2715), caudal vertebra (AMNH 2702) and femur (AMNH 2704). Cope's diagnosis consists of characters found in all coelophysoids (cervical centra with concave anterior articular surface; oblique articular surfaces on cervical centra; slender mid caudal vertebrae), as well as the supposed absence of posterior pleurocoels in the cervical (untrue), and the larger size than C. bauri (possibly ontogenetic or individual variation). Cope later (1887b) reassigned the species to Tanystropheus due to the amphicoelous cervicals and referred several more specimens (AMNH 2703, 2705-2708, 2716, 2735). Huene (1906) further referred a distal metapodial (AMNH 2730) to the species, and later (1915) reassigned the material quite extensively. The syntype dorsal (AMNH 2715) was referred to C. bauri, as was the ilium AMNH 2708. The caudal AMNH 2735 was reassigned to either C. bauri or C. willistoni, while the supposed tibia or metatarsal AMNH 2721 that had been referred to C. bauri was now referred to C. longicollis (it's actually a Dromomeron femur- Nesbitt et al., 2009). Huene (1915) also referred several additional specimens (AMNH 2729, 2731, 2739, 2749) to C. longicollis. Though Welles (1984) intended his Longosaurus longicollis to be based on this species, he chose a holotype different than the C. longicollis' lectotype, so the taxa are not objective synonyms. See the discussion of "unnamed Coelophysidae (Cope, 1887)" above for more details. The lectotype has an elongate centrum as in coelophysids and shows the anterior pedicular foramina which are apomorphic for Coelophysis bauri. It is thus synonymized with that species here.
Ghost Ranch material- In 1947 an extensive bonebed of coelophysoids was discovered at the Ghost Ranch quarry (also called the Whitaker quarry) and were assigned to Coelophysis by Colbert (1947) in a popular article. Colbert and Baird later (1958) assigned the Ghost Ranch specimens specifically to C. bauri, which Colbert (1964) explained was due to his believing C. longicollis and C. willistoni were older and younger individuals respectively of the same species. Hunt and Lucas (1991) felt Cope's original Coelophysis specimens (including the lectotypes of C. bauri, C. longicollis and C. willistoni) were indeterminate, and created the name Rioarribasaurus colberti for the abundant complete Ghost Ranch specimens. However, the ICZN later (1996) ruled a Ghost Ranch specimen is the neotype of Coelophysis bauri. Coelophysis? longicollis, C? willistoni, the other original Coelophysis specimens, Longosaurus, Podokesaurus and Protoavis (in part) may all be referrable to C. bauri or at least Coelophysis, but this cannot be determined until the Ghost Ranch specimens are redescribed. The illustrations of the latter in Colbert (1989) are inaccurate and have hampered comparison to other coelophysids. Downs (2000) notes most of the supposed differences from Megapnosaurus rhodesiensis are not real (C. bauri has a nasal fenestra; vaulted palate; interdental plates; obturator fenestra; pubic fenestra; fused pelvis; triangular dorsal transverse processes; identical hindlimb morphology). Tykoski (2005) notes the presence of a median basisphenoid spur that follows the roof of the basisphenoid recess cannot be ascertained in Coelophysis (contra Tykoski and Rowe, 2004). Ezcurra (2006) noted that contra Bristowe and Raath (2004), an anteriorly pointed antorbital fossa only occurs in some adult specimens (e.g. CM C-3-82, AMNH 7224, YPM 41196), but not others (e.g. AMNH 7240, MCZ 4327). It is thus not a diagnostic character of the species.
Snyder Quarry material- The Snyder Quarry coelophysoid material (NMMNH P-29046-29047, 30779-30780, 30852, 31293-31661) was originally referred to Eucoelophysis sp. by Heckert et al. (2000) based on supposed similarities in the scapulocoracoid, ischium and tibia. Nesbitt et al. (2007) showed the former genus is non-dinosaurian while Heckert et al.'s material exhibits numerous dinosaur, theropod and coelophysoid synapomorphies. Eucoelophysis' holotype differs in lacking a distinct acetabular rim on the ischium, an offset femoral head, a differentiated lateral and fibular condyle that is separated by a distinct sulcus, and a well developed curved cnemial crest. The appressed lateral tibial surface is developed differently in both taxa. Ezcurra (2006) noted NMMNH P-30852 shows an alveolar ridge like Liliensternus and more derived coelophysoids, and a square anterior end on the antorbital fossa as in Zupaysaurus and more derived coelophysoids. Spielmann et al. (2007) redescribed the material (including newly identified NMMNH P-54617-54620), assigning it to Coelophysis bauri based on NMMNH P-30852 lacking a promaxillary fenestra (also in Megapnosaurus) and having a ventrally broad lacrimal. The other Snyder Quarry material was assigned to the same species based on general similarity and close association, though the other specimens were only identified to more inclusive taxa based on their own morphologies. While Spielmann et al. identified NMMNH P-54618-54620 as Coelophysis based on the transverse proximal femoral groove, this is also present in Kayentavenator.
Cannibalism in Coelophysis?- AMNH 7223 and 7224 are preserved with supposed stomach contents (vertebrae and a hindlimb in 7223; articulated remains anteriorly, plus a sacral vertebra, ilium and proximal femora posteriorly in 7224) that have been traditionally viewed as evidence of cannibalism (e.g. Colbert, 1989). However, the supposed stomach contents of AMNH 7223 don't lie within the ribcage, and those in the anterior of 7224 lie under the ribcage (Gay, 2002; Nesbitt et al., 2006). The posterior fragments in AMNH 7224 are within the ribcage, but are crocodylomorph, not theropod (Nesbitt et al., 2006). The supposedly cannibalized manual elements identified in coprolites and cololites by Rinehart et al. (2005) cannot be identified as Coelophysis (Nesbitt et al., 2006). There is thus no evidence of cannibalism in Coelophysis.
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Rinehart, Hunt, Lucas and Heckert, 2005. Coprolites and cololites from the Late Triassic theropod dinosaur Coelophysis bauri, Whitaker Quarry, Rio Arriba County, NM. New Mexico Geology. 27(2), 53.
Rinehart, Hunt, Lucas, Heckert and Smith, 2005. New evidence of cannibalism in the Late Triassic (Apachean) dinosaur, Coelophysis bauri (Theropoda: Ceratosauria). Journal of Vertebrate Paleontology. 25(3), 105A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas. 28(4), 649-684.
Nesbitt, Turner, Erickson and Norell, 2006. Prey choice and cannibalistic behaviour in the theropod Coelophysis. Biology Letters. 2(4), 611-614.
Rinehart, Lucas and Hunt, 2006. The furcula of Coelophysis bauri, a Late Triassic (Apachean) dinosaur (Theropoda: Ceratosauria) from New Mexico. New Mexico Geology. 28(2), 62.
Ezcurra, 2007. The cranial anatomy of the coelophysoid theropod Zupaysaurus rougieri from the Upper Triassic of Argentina. Historical Biology. 19(2), 185-202.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.
Spielmann, Lucas, Rinehart, Hunt, Heckert and Sullivan, 2007. Oldest records of the Late Triassic theropod dinosaur Coelophysis bauri. New Mexico Museum of Natural History and Science Bulletin. 41, 384-401.
Rinehart, Lucas, Heckert, Spielmann and Celeskey, 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, 260 pp.
Gay, 2010. Notes on Early Mesozoic theropods. Lulu Press. 44 pp.

Megapnosaurus Ivie, Slipinski and Wegrzynowicz, 2001
= Syntarsus Raath, 1969 (preoccupied Fairmaire, 1869)
Comments- This genus is often synonymized with Coelophysis (Paul, 1988; Bristowe and Raath, 2004), as new discoveries invalidate supposed differences between the taxa (e.g. Downs, 2000; Bristowe and Raath, 2004). In 2001, it was discovered that Syntarsus was preoccupied by a beetle. The entomologists who determined this attempted unsuccessfully to contact Raath so that he could rename it, and ended up renaming it themselves (Ivie et al., 2001). Paleontologists might have reacted more positively if the replacement name (Megapnosaurus) hadn't been facetious, translating to "big dead lizard", since that's what all dinosaurs are to entomologists. Thus the trend has been to ignore the name, a position made easier by its resemblence to Coelophysis. However, personal dislike for a name is not a valid reason to reject it, and Coelophysis and Megapnosaurus are more closely related to each other than either is to other well known coelophysoids (Procompsognathus, Segisaurus, Liliensternus, Gojirasaurus, etc.). Thus their synonymy is a subjective decision, which some workers follow (Downs, 2000; Bristowe and Raath, 2004) and others do not (Irimis, 2004; Tykoski and Rowe, 2004). I keep them separate primarily due to the influence of Tykoski and Rowe's paper.
M. rhodesiensis (Raath, 1969) Ivie, Slipinski and Wegrzynowicz, 2001
= Syntarsus rhodesiensis Raath, 1969
= Coelophysis rhodesiensis (Raath, 1969) Paul, 1988
Hettangian-Sinemurian, Early Jurassic
Upper Elliot Formation, South Africa; Forest Sandstone Formation, Zimbabwe
Holotype- (QG 1) (2.15 m, 13 kg) (robust adult) (skull ~210 mm), fifth dorsal vertebra (28 mm), sixth dorsal vertebra (28 mm), seventh dorsal vertebra (29 mm), eighth dorsal vertebra (29 mm), ninth dorsal vertebra (29 mm), tenth dorsal vertebra (32 mm), eleventh dorsal vertebra (31 mm), twelfth dorsal vertebra (29 mm), thirteenth dorsal vertebra (29 mm), twelve dorsal ribs (sixth 141 mm), gastralia, sacrum (123 mm), first caudal vertebra (24 mm), second-ninth caudal vertebrae, tenth caudal vertebra (27 mm), eleventh to nineteenth caudal vertebra, twenty-second caudal vertebra (26 mm), twenty-third to twenty-fifth caudal vertebra, twenty-sixth caudal vertebra (27 mm), twenty-seventh caudal vertebra (29 mm), twenty-eighth caudal vertebra (31 mm), twenty-ninth caudal vertebra, thirtieth caudal vertebra (26 mm), thirty-first to thirty-eighth caudal vertebra, thirty-ninth caudal vertebra (22 mm), fortieth caudal vertebra, thirty-six chevrons, scapulocoracoid (130 mm), humerus (100 mm), radius (61 mm), ulna (77 mm), six carpals, metacarpal I (12 mm), phalanx I-1 (20 mm), manual ungual I (18 mm), metacarpal II (26 mm), phalanx II-1 (13 mm), phalanx II-2 (17 mm), manual ungual II (19 mm), metacarpal III (26 mm), phalanx III-1 (10 mm), phalanx III-2 (9 mm), phalanx III-3 (12 mm), manual ungual III, metacarpal IV (17 mm), phalanx IV-1 (5 mm), ilium (148 mm), pubis (204 mm), ischium (130 mm), femur (208 mm), tibia (223 mm), fibula (208 mm), astragalocalcaneum (29 mm wide), distal tarsal IV, metatarsal I (30 mm), phalanx I-1 (17 mm), pedal ungual I (14 mm), metatarsal II (119 mm), phalanx II-1 (33 mm), phalanx II-2 (25 mm), pedal ungual II (22 mm), metatarsal III (132 mm), phalanx III-1 (37 mm), phalanx III-2 (29 mm), phalanx III-3 (24 mm), pedal ungual III (22 mm), metatarsal IV (117 mm), phalanx IV-1 (20 mm), phalanx IV-2 (17 mm), phalanx IV-3 (15 mm), phalanx IV-4 (12 mm), pedal ungual IV (18 mm), metatarsal V (47 mm)
Referred- (BPI coll.) (at least eight individuals) cervical vertebra, dorsal vertebra, distal femora, proximal tibia, pedal fragments (Raath 1980)
(QG 3A) (robust) femur (Raath, 1990)
(QG 76) (gracile) femur (201 mm) (Raath, 1990)
(QG 164) pes (Raath, 1977)
(QG 165) maxilla, nasals, lacrimal, jugal, quadratojugal, palatine, ectopterygoid, pterygoid, hyoids, postcrania (Bristowe and Raath, 2004)
(QG 169) seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth cervical vertebra, first dorsal vertebra, second dorsal vertebra, third dorsal vertebra, fourth dorsal vertebra (Raath, 1977)
(QG 170) eighth cervical vertebra (Raath, 1977)
(QG 171) second dorsal vertebra (Raath, 1977)
(QG 172) sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra (Raath, 1977)
(QG 173) cervical ribs (Raath, 1977)
(QG 175) third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra (Raath, 1977)
(QG 176) atlas, axis, third cervical vertebra (Raath, 1977)
(QG 177) second cervical vertebra, third cervical vertebra, fourth cervical vertebra (Raath, 1977)
(QG 179) third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra (Raath, 1977)
(QG 180) second dorsal vertebra (Raath, 1977)
(QG 193) maxilla, nasal, lacrimal, prefrontal, jugal, frontals, parietals, postorbital, squamosal, quadratojugal, braincase, mandible, scapulocoracoid, two furculae (Raath, 1977; Tykoski et al., 2002)
(QG 194) maxilla, nasals, squamosal, quadratojugal, quadrate, braincase, endocranium, ectopterygoid, pterygoid (Raath, 1977)
(QG 195) braincase, endocranium (Raath, 1977)
(QG 196) braincase, endocranium (Raath, 1977)
(QG 197) braincase, endocranium (Raath, 1977)
(QG 202) premaxilla, maxilla, nasal, dentaries, dentary teeth (Raath, 1977)
(QG 206) maxilla (Raath, 1977)
(QG 208) maxilla, sclerotic plates, mandible (Raath, 1977)
(QG 209) maxilla (Raath, 1977)
(QG 210) maxilla (Raath, 1977)
(QG 213) maxilla (Raath, 1977)
(QG 230) lacrimal (Raath, 1977)
(QG 234) lacrimal (Raath, 1977)
(QG 235) quadratojugal, quadrate, palatine, ectopterygoid, pterygoid (Raath, 1977)
(QG 241) palatine (Raath, 1977)
(QG 244) furcula (Raath, 1977; Tykoski et al., 2002)
(QG 245) premaxilla, premaxillary teeth (Raath, 1977)
(QG 246) premaxilla, premaxillary teeth (Raath, 1977)
(QG 249) premaxilla (Raath, 1977)
(QG 254) premaxilla (Raath, 1977)
(QG 263) pterygoid (Raath, 1977)
(QG 265) pterygoid (Raath, 1977)
(QG 278) jugal, lacrimal, prefrontal (Raath, 1977)
(QG 287) postorbital (Raath, 1977)
(QG 303) dentary (Raath, 1977)
(QG 305) dentary (Raath, 1977)
(QG 307) mandible (Raath, 1977)
(QG 406) fifth dorsal vertebra (Raath, 1977)
(QG 408) seventh dorsal vertebra (Raath, 1977)
(QG 514) (robust) humerus (Raath, 1990)
(QG 514b) (robust) ulna (Raath, 1990)
(QG 545) (gracile) humerus (Raath, 1990)
(QG 563) ulna
(QG 568) (gracile) ulna (Raath, 1990)
(QG 573) (robust) metacarpus (Raath, 1990)
(QG 577) (gracile) distal radius, distal ulna, manus (Raath, 1990)
(QG 687) distal carpal (Raath, 1977)
(QG 691) (gracile) pelvis, femur (142 mm), tibia, fibula, astragalus, calcaneum (Raath, 1990)
(QG 713) (gracile) femur (Raath, 1990)
(QG 715) (gracile) femur (Raath, 1990)
(QG 716) femur (Raath, 1990)
(QG 717) (gracile) femur (Raath, 1990)
(QG 725) (robust) femur (Raath, 1990)
(QG 726) (robust) femur (Raath, 1990)
(QG 727) (robust) femur (Raath, 1990)
(QG 729) (robust) femur (Raath, 1990)
(QG 731) (robust) femur (189 mm) (Raath, 1990)
(QG 732) (robust) femur (Raath, 1990)
(QG 733) (robust) femur (Raath, 1990)
(QG 738) (gracile) femur (192 mm) (Raath, 1990)
(QG 739) (gracile) femur (Raath, 1990)
(QG 740) (gracile) femur (Raath, 1990)
(QG 742) (gracile) femur (Raath, 1990)
(QG 744) (gracile) femur (Raath, 1990)
(QG 745) (gracile) femur (Raath, 1990)
(QG 753) (robust) femur (Raath, 1990)
(QG 754) (robust) femur (186 mm) (Raath, 1990)
(QG 755) (robust) femur (185 mm) (Raath, 1990)
(QG 760) (robust) femur (Raath, 1990)
(QG 768) (gracile) tibia, fibula, astragalus, calcaneum, distal tarsal III, distal tarsal IV, metatarsus (Raath, 1990)
(QG 786) (gracile) astragalus, calcaneum (Raath, 1990)
(QG 816) astragalus, calcaneum (Raath, 1977)
(QG 846) distal tarsal III (Raath, 1977)
Diagnosis- (modified from Tykoski and Rowe, 2004) differs from Coelophysis bauri in the pit at the base of the nasal process of the premaxilla; Differs from "Megapnosaurus" kayentakatae in promaxillary fenestra absent; nasal crests absent; frontals not separated by midline anterior extension of parietals; anterior astragalar surface flat.
(after Ezcurra, 2006) reduced distal medial condyle on metacarpal I.
(after Carrano et al., 2012) blunt, squared anterior margin of antorbital fossa; base of lacrimal vertical ramus width <30% its height; maxillary and dentary tooth rows end posteriorly at anterior rim of lacrimal
Comments- Munyikwa and Raath (1999) described a partial skull as Syntarsus, which was tentatively reidentified as Dracovenator by Yates (2005). Bristowe and Raath (2004) used a partially articulated juvenile skull to show the nasal fenestra identified by Raath (1977) was in fact closed in life, Raath articulated the palatine backwards, and that his reconstruction of the lacrimal-jugal articulation is inaccurate. They also confirmed the hyoids identified by Raath were furculae.
References- Raath, 1969. A new Coelurosaurian dinosaur from the Forest Sandstone of Rhodesia. Arnoldia Rhodesia. 4 (28): 1-25.
Raath, 1977. The anatomy of the Triassic theropod Syntarsus rhodesiensis (Saurischia: Podokesauridae) and a consideration of its biology. PhD thesis. Rhodes University.
Raath, 1980. The theropod dinosaur Syntarsus (Saurischia: Podokesauridae) discovered in South Africa. S. Afr. J. Sci. 76: 375-376.
Raath, 1985. The theropod Syntarsus and its bearing on the originn of birds. p. 219-227 in Max K. Hecht, et al. (eds.), The beginnings of birds. Freunde des Jura-Museums Eichstätt, Eichstätt.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Raath, 1990. Morphological variation in small theropods and its meaning in systematics: Evidence from Syntarsus rhodesiensis. p. 91-105 in Kenneth Carpenter, Philip J. Currie (eds.), Dinosaur systematics. Approaches and perspectives. Cambridge University Press, Cambridge, etc..
Chinsamy, 1990. Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda). Palaeontol. Afr. 27 p. 77-82.
Munyikwa and Raath, 1999. Further material of the ceratosaurian dinosaur Syntarsus from the Elliot Formation (Early Jurassic) of South Africa. Palaeontologia Africana. 35:55-59.
Ivie, Slipinski and Wegrzynowicz, 2001. Generic homonyms in the Colydiinae (Coleoptera: Zopheridae). Insecta Mundi. 15, 63-64.
Tykoski, Forster, Rowe, Sampson and Munyikwa, 2002. A furcula in the coelophysid theropod Syntarsus. Journal of Vertebrate Paleontology. 22(3):728–733.
Bristowe and Raath, 2004. A juvenile coelophysoid skull from the Early Jurassic of Zimbabwe, and the synonymy of Coelophysis and Syntarsus. Palaeontologia Africana. 40, 31-41.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods. Palaeontologia Africana. 41, 105-122.
Ezcurra, 2007. The cranial anatomy of the coelophysoid theropod Zupaysaurus rougieri from the Upper Triassic of Argentina. Historical Biology. 19(2), 185-202.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.