Avepoda Paul, 2002
Definition- (metatarsal I does not contact distal tarsals homologous with Allosaurus fragilis) (modified from Paul, 2002)
= Paleotheropoda Paul, 1988
Comments- This clade was first suggested by Paul (1988) as a more appropriate name for Theropoda (as their feet are birdlike as opposed to beastlike), though he recognized Theropoda could not be renamed. In his later 2002 book, Paul actually proposed Avepoda as a subgroup of Theropoda, for those "that either possessed a foot in which metatarsal I did not contact the distal tarsals, or descended from such theropods, and belonged to a clade which includes Neotheropoda." The definition is here modified by using Allosaurus fragilis as the internal specifier, as that taxon is the official internal specifier of Theropoda. Avepoda has almost exclusively been used by Paul, with other authors using Neotheropoda (sensu Sereno) for the group since the two usually have the same known content (though Avepoda is near certainly more inclusive than Neotheropoda sensu Sereno, unless coelophysoids were the first theropods to develop the tridactyl pes). However, Procompsognathus was an avepod possibly outside the Coelophysis+Passer clade in Paul's (1988) phylogeny and Gauthier's (1986) analysis, as was Liliensternus in the latter. More recently, Liliensternus was found to be outside the Coelophysis+Passer clade in Bittencourt Rodrigues' (2010) thesis. A different complication arises in Martinez et al.'s (2011) cladogram, where the tetradactyl Tawa is a basal coelophysoid. In this case, Avepoda contains Tawa and other coelophysoids under ACCTRAN character optimization, but excludes Coelophysoidea under DELTRAN optimization. Note it is not the same as Avipoda Novas, 1992, which was proposed for a clade similar in extent to Tetanurae.
Paleotheropoda was proposed as a paraphyletic order of theropods by Paul (1988), including all theropods which were not avetheropods. This consisted of what are generally recognized today as coelophysoids, ceratosaurs, megalosauroids and sinraptorids. No other reference used the term due to the general dislike of paraphyletic groups, and in 2002 Paul referred to the same grade as baso-avepods. As Paul's (1988) Theropoda was equivalent to Avepoda, Paleotheropoda is catalogued here as a synonym of that group.
References- Gauthier, 1986. Saurischian monophyly and the origin of birds. Memoirs of the Californian Academy of Sciences 8, 1-55.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster, New York. 464 pp
Paul, 2002. Dinosaurs of the Air. The John Hopkins University Press, Baltimore and London. 460 pp.
Bittencourt Rodrigues, 2010. Revisao filogenetica dos dinossauriformes basais: Implicacoes para a origem dod dinossauros. Unpublished Doctoral Thesis. Universidade de Sao Paulo. 288 pp.
Martinez, Sereno, Alcober, Columbi, Renne, Montanez and Currie, 2011. A basal dinosaur from the dawn of the dinosaur era in southwestern Pangaea. Science. 331, 206-210.

undescribed neotheropod (Britt, Chure, Engelemann, Scheetz and Hansen, 2010)
Late Norian-Rhaetian, Late Triassic
Saints and Sinners Quarry, Nugget Sandstone, Utah, US
Material- teeth (~30 mm)
References- Britt, Chure, Engelemann, Scheetz and Hansen, 2010. Multi-taxic theropod bonebeds in an interdunal setting of the Early Jurassic Eolian Nugget Sandstone, Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 65A.
Britt, Chambers, Engelmann, Chure and Scheetz, 2011. Taphonomy of coelophysoid bonebeds preserved along the shoreline of an Early Jurassic lake in the Nugget Sandstone of NE Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 78.

undescribed Neotheropoda (Kirkland and Milner, 2005)
Hettangian, Early Jurassic
Freeman Quarry SGDS14V, Whitmore Point Member of the Moenave Formation, Utah, US
Material- (SGDS 851) tooth (Milner and Lockley, 2006)
(SGDS 852) tooth (Milner and Lockley, 2006)
(SGDS 1335) incomplete tooth ( Milner, Birthisel, Kirkland, Breithaupt, Matthews, Lockley, Santucci, Gibson, DeBlieux, Hurlbut, Harris and Olsen, 2012)
(SGDS coll.) several teeth (Milner and Lockley, 2006)
Comments- Kirkland and Milner (2005) first mention "Bones and teeth of theropods large enough to produce Eubrontes tracks" from the Whitmore Point Member, while Milner and Lockley (2006) note specifically "a possible Megapnosaurus tooth (Fig. 7G) from "Freeman Quarry"", which was figured by Milner et al. (2012) as "Small serrated tooth, possibly from a coelophysoid theropod (SGDS 851)." Milner and Lockley also mention "several of the well-preserved dinosaur teeth found in Freeman Quarry on WCSD property (Figure 7I) resemble those of the early Cretaceous fish-eating dinosaur, Spinosaurus" and figure a specimen identified as SGDS 852 in Milner et al.. They state these teeth "are conical and sometimes preserve serrations under the right circumstances" but are careful to note "they are not spinosaurids", and figure an additional example.
References- Kirkland and Milner, 2005. The case for theropod dinosaurs exploiting fish as a major food resource during the Early Jurassic. Tracking Dinosaur Origins: The Triassic/Jurassic Terrestrial Transition Abstracts Volume. 9-10.
Milner and Lockley, 2006. History, geology, and paleontology: St. George Dinosaur Discovery Site at Johnson Farm, Utah. In Reynolds (ed.). Making Tracks Across the Southwest, Abstracts from the 2006 Desert Symposium. 35-48.
Milner, Birthisel, Kirkland, Breithaupt, Matthews, Lockley, Santucci, Gibson, DeBlieux, Hurlbut, Harris and Olsen, 2012. Tracking Early Jurassic dinosaurs across southwestern Utah and the Triassic-Jurassic transition. Nevada State Museum Paleontological Papers. 1, 1-107.

unnamed Neotheropoda (Hunt, 1998)
Early Norian, Late Triassic
Dinosaur Wash, Dying Grounds PFV 122, Blue Mesa Member of Chinle Formation, Arizona, US
?(MDM coll.) cranial fragments, teeth, vertebrae (Hunt and Wright, 1999)
Early Norian, Late Triassic
Dinosaur Ridge PFV 211, Blue Mesa Member of Chinle Formation, Arizona, US
(PEFO 43909; = NMMNH coll.; = MDM coll.) proximal tibia (Hunt, 1998)
Early Norian, Late Triassic
North Stinking Springs SMU 252, Blue Mesa Member of Chinle Formation, Arizona, US
?(SMU coll.) teeth, vertebrae (Polcyn, Winkler, Jacobs and Newman, 2002)
Comments- One of two specimens discovered in 1996 at the Dinosaur Ridge locality and assigned by Hunt (1998) to Theropoda, he notes "The larger species is only known at present from a proximal tibia that represents an animal comparable in size to the specimen described from Norian strata in PEFO by Padian (1986)." Nesbitt et al. (2007) stated "The proximal portion of the tibia bears a cnemial crest and two divided posterior condyles. These features are not diagnostic to Theropoda; thus, this specimen cannot be assigned to the Theropoda" and so only assigned it to Archosauria indet.. While Hunt and Wright list it as in the MDM coll. and Nesbitt et al. as NMMNH unnumbered, Marsh and Parker (2020) reveal it eventually became PEFO 43909. Marsh and Parker assigned it to Neotheropoda after all based on the concave surface between posterior condyles and cnemial crest and the fibular crest.
In 1998 the Dinosaur Wash part of the Dying Grounds locality was discovered, and Hunt and Wright (1999) state it "yields diverse small reptile specimens including abundant postcrania and teeth of a small theropod" depopsited at the MDM. Therrien and Fastovsky (2000) state "The theropod remains consist of vertebrae, teeth, and skull fragments belonging to an undetermined theropod."
Polcyn et al. (2002) state "ceratosaur vertebral elements and teeth that are probably referrable to the Coelophysidae" are present at SMU microvertebrate locality 252, listed as ?Coelophysis sp. in their faunal table. Parker (2005) says "obviously these may also belong to poposaurids as admitted by these workers", but Polcyn et al. only said this about different "small, laterally compressed teeth [that] may be referable to either theropods or poposaurs." Parker similarly misrepresented the paper when he claimed "these authors note that several jaw fragments with teeth may actually pertain to coelophysoids", when Polcyn et al. actually wrote "numerous isolated teeth and tooth-bearing jaw fragments have been recovered and are referable to sphenodontids, lepidosauromorphs, and theropod and ornithischian dinosaurs" without explicitly saying any of the jaw fragments were theropod. In any case the material remains undescribed, and while isolated teeth are usually best referred to Archosauria indet., vertebrae could be more diagnostic if they are more than dorsal or caudal centra.
References- Hunt, 1998. Preliminary results of the Dawn of the Dinosaurs Project Petrified Forest National Park, Arizona. In Santucci and McClelland (eds.). National Park Service Paleontological Research. National Park Service Technical Report NPS/NRGRD/GRDTR-98/1. 135-137.
Hunt and Wright, 1999. New discoveries of Late Triassic dinosaurs from Petrified Forest National Park, Arizona. in Santucci and McClelland (eds.). National Park Service Paleontological Research Volume 4. Geologic Resources Division Technical Report NPS/NRGRD/GRDTR-99/03. 96-100.
Therrien and Fastovsky, 2000. Paleoenvironments of early theropods, Chinle Formation (Late Triassic), Petrified Forest National Park, Arizona. PALAIOS. 15(3), 194-211.
Polcyn, Winkler, Jacobs and Newman, 2002. Fossil occurrences and structural disturbance in the Triassic Chinle Formation at North Stinking Springs Mountain near St. Johns, Arizona. New Mexico Museum of Natural History and Science Bulletin. 21, 43-49.
Parker, 2005. Faunal review of the Upper Triassic Chinle Formation of Arizona. In McCord (ed.). Vertebrate Paleontology of Arizona. Mesa Southwest Museum Bulletin. 11, 34-54.
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.
Marsh and Parker, 2020. New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils. PaleoBios. 37, 1-56.

unnamed Neotheropoda (Tannenbaum, 1983)
Middle Norian, Late Triassic
Bowman Site PFV 089, Sonsela Member of the Chinle Formation, Arizona, US
Material- (PEFO 31187) proximal femur (Parker and Irmis, 2005)
Middle Norian, Late Triassic
Kaye Quarry PFV 410, Sonsela Member of the Chinle Formation, Arizona, US
(PEFO 39421; = UWBM 108881) proximal tibia (Marsh and Parker, 2020)
(PEFO 39563; = UWBM 109902) proximal femur (Marsh and Parker, 2020)
Middle Norian, Late Triassic
Placerias Quarry UCMP A269, Sonsela Member of the Chinle Formation, Arizona, US
?(MNA coll.) (Tannenbaum, 1983)
(UCMP 25820) distal tibia (Irmis, 2005)
Comments- PEFO 31187 was stated by Parker and Irmis (2005) to "possess a sloping posterior margin adjacent to the greater trochanter and therefore [is] probably referable to the Theropoda." All of the PEFO specimens were assigned to Neotheropoda by Marsh and Parker (2020) based on "An enlarged anteromedial tuber and corresponding ligament sulcus" or "the presence of the fibular crest and a concave proximal surface between the cnemial crest and posterior condyles."
Discovered in 1934, Irmis (2005) stated UCMP 25820 is "assignable to the Saurischia because it has a well-developed posterolateral process, is quadrangular in distal view, and has a concave posterolateral margin in distal view." Nesbitt et al. (2007) instead stated the presence of "a well-developed descending posterolateral process, a concave posterolateral margin in distal view and a well developed dorsal excavation for insertion of the ascending process of the astragalus," ... "in combination with a posterolateral process that extends well beyond the body of the tibia laterally, allow us to refer this specimen to Theropoda indet." They state it may be referrable to Camposaurus, which should be determinable based on the tibial autapomorphies of that genus.
Tannenbaum (1983) references "an unnumbered and undescribed theropod specimen at the Museum of Northern Arizona in the macrofaunal collection from the Placerias Quarry (R. Long, pers. comm.)."
References- Tannenbaum, 1983. The microvertebrate fauna of the Placerias and Downs' quarries, Chinle Formation (Upper Triassic) near St. Johns, Arizona. Masters thesis, University of California. 111 pp.
Irmis, 2005. The vertebrate fauna of the Upper Triassic Chinle Formation in northern Arizona. In Nesbitt, Parker and Irmis (eds.). Guidebook to the Triassic Formations of the Colorado Plateau in Northern Arizona: Geology, Paleontology, and History. Mesa Southwest Museum, Bulletin. 9, 63-88.
Parker and Irmis, 2005. Advances in Late Triassic vertebrate paleontology based on new material from Petrified Forest National Park, Arizona. New Mexico Museum of Natural History and Science Bulletin. 29, 45-58.
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.
Marsh and Parker, 2020. New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils. PaleoBios. 37, 1-56.

unnamed Neotheropoda (Hunt, Olson, Huber, Shipman, Bircheff and Frost, 1996)
Late Norian, Late Triassic
Dinosaur Hill PFV 040 / Inadvertent Hills UCMP V82250, Petrified Forest Member of Chinle Formation, Arizona, US
Material- (PEFO coll.) vertebrae, femur (Hunt, Olson, Huber, Shipman, Bircheff and Frost, 1996)
Late Norian, Late Triassic
Flattops, Petrified Forest Member of the Chinle Formation, Arizona, US
Material- ?(MDM coll.) limb element shaft (Hunt and Wright, 1999)
Late Norian, Late Triassic
Zuni Well Mound PFV 215, Petrified Forest Member of the Chinle Formation, Arizona, US
?(MDM coll.) vertebrae and/or limb fragments (Hunt and Wright, 1999)
Late Norian, Late Triassic
RAP Hill PFV 216, Petrified Forest Member of the Chinle Formation, Arizona, US
?(MDM coll.) vertebrae and/or limb fragments (Hunt and Wright, 1999)
Late Norian, Late Triassic
The Giving Site PFV 231, Petrified Forest Member of the Chinle Formation, Arizona, US
(PEFO 33984) proximal femur (Parker and Irmis, 2005)
(PEFO 34079) proximal femur (Marsh and Parker, 2020)
(PEFO 34080) proximal tibia (Marsh and Parker, 2020)
(PEFO 34613) proximal femur (Marsh and Parker, 2020)
Late Norian, Late Triassic
RAP Hill North PFV 277, Petrified Forest Member of the Chinle Formation, Arizona, US
?(MDM coll.) vertebrae and/or limb fragments (Hunt and Wright, 1999)
Late Norian, Late Triassic
Rabbit Foot Hills PFV 302, Petrified Forest Member of the Chinle Formation, Arizona, US
(PEFO 38714) proximal femur (Marsh and Parker, 2020)
Late Norian, Late Triassic
Black Knoll E PFV 451, Petrified Forest Member of the Chinle Formation, Arizona, US
(PEFO 44468) proximal tibia (Marsh and Parker, 2020)
Late Norian, Late Triassic
Sorrel Horse Mesa SE PFV 475, Petrified Forest Member of the Chinle Formation, Arizona, US
(PEFO 43506) proximal femur (Marsh and Parker, 2020)
(PEFO 43550) proximal femur (Marsh and Parker, 2020)
(PEFO 44472) proximal femur (Marsh and Parker, 2020)
Late Norian, Late Triassic
The Corner PFV 477, Petrified Forest Member of the Chinle Formation, Arizona, US
(PEFO 44473) proximal femur (Marsh and Parker, 2020)
Late Norian, Late Triassic
RAP Hill West, Petrified Forest Member of the Chinle Formation, Arizona, US
?(MDM coll.) vertebrae and/or limb fragments (Hunt and Wright, 1999)
Comments- Hunt et al. (1996) first mention "undescribed specimens of a smaller theropod dinosaur" and "specimens (vertebrae, femur) of a ceratosaurian distinctly smaller than the "Coelophysis" partial skeleton." Hunt (1998) wrote "Our collecting has yielded specimens of a third, smaller, ?ceratosaurian dinosaur including a complete femur and numerous vertebra" from the Dinosaur Hill locality", while Hunt et al. (1998) stated "Several other theropod specimens (femur, vertebrae) have recently been collected by APH from this locality (uncatalogued PEFO specimens)." These would have been collected in 1996 as part of The Dawn of the Dinosaurs Project and used Hunt's sensu lato concept of Ceratosauria that includes coelophysoid-grade taxa, but have yet to be described.
Hunt and Wright (1999) mentioned "a hollow, theropod limb shaft from Flattops" collected in 1998 by the MDM in the 'Painted Desert' (= Petrified Forest) Member. This locality might be Flattops NW (PFV 070) or Flattops W (PFV 071) (but is not Flattops NE PFV 354 because that is in the Sonsela Member), and the specimen is possibly Archosauria indet. in any case if even the identity of the limb bone can't be determined. They also say "The second most common dinosaur[in the Petrified Forest Member] is a small theropod that is represented at most localities by vertebrae and limb fragments", and their Table 1 lists MDM material found in 1998 at Zuni Well Mound and RAP Hill, as well as MDM material found in 1999 at RAP Hill North and RAP Hill West. Marsh and Parker (2020) describe a proximal femur (PEFO 34863) and metatarsal I (PEFO 36741) from Zuni Well Mound which they refer to Dinosauria and Ornithodira respectively, which may be some of the supposed theropod limb fragments from that locality.
PEFO 33984 was stated by Parker and Irmis (2005) to "possess a sloping posterior margin adjacent to the greater trochanter and therefore [is] probably referable to the Theropoda." All of the PEFO specimens were assigned to Neotheropoda by Marsh and Parker (2020) based on "An enlarged anteromedial tuber and corresponding ligament sulcus" or "the presence of the fibular crest and a concave proximal surface between the cnemial crest and posterior condyles."
References- Hunt, Olson, Huber, Shipman, Bircheff and Frost, 1996. A new theropod locality at Petrified Forest National Park with a review of Late Triassic dinosaur localities in the park. Fossils of Arizona Symposium, 4, 55-61.
Hunt, 1998. Preliminary results of the Dawn of the Dinosaurs Project Petrified Forest National Park, Arizona. In Santucci and McClelland (eds.). National Park Service Paleontological Research. National Park Service Technical Report NPS/NRGRD/GRDTR-98/1. 135-137.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the western United States. Geobios. 31(4), 511-531.
Hunt and Wright, 1999. New discoveries of Late Triassic dinosaurs from Petrified Forest National Park, Arizona. In Santucci and McClelland (eds.). National Park Service Paleontological Research Volume 4. Geologic Resources Division Technical Report NPS/NRGRD/GRDTR-99/03. 96-100.
Parker and Irmis, 2005. Advances in Late Triassic vertebrate paleontology based on new material from Petrified Forest National Park, Arizona. New Mexico Museum of Natural History and Science Bulletin. 29, 45-58.
Marsh and Parker, 2020. New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils. PaleoBios. 37, 1-56.

unnamed neotheropod (Kirby, 1991)
Rhaetian, Late Triassic
MNA 853, Owl Rock Member of the Chinle Formation, Arizona, US
Material- (MNA.V.7240) proximal femur
Comments- This was assigned to Ceratosauria? indet. by Kirby (1991), who described it and stated it "is less anteroposteriorly flattened than in Syntarsus, but corresponds closely to the robust trochanter condition in Coelophysis." Hunt et al. (1998) mentioned it as undoubtedly dinosaurian, and Spielmann et al. (2007) found it was labeled Coelophysis sp. in the MNA collections and used the ""hooked" femoral head and a prominent trochanteric shelf" to refer it to Coelophysoidea. Given the recent interpretation of coelophysoids as a basal grade of neotheropods and similarity to e.g. Liliensternus, it is assigned to Neotheropoda here.
References- Kirby, 1991. A vertebrate fauna from the Upper Triassic Owl Rock Member of the Chinle Formation of northern Arizona. Masters thesis, Northern Arizona University. 496 pp.
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.). New Mexico Museum of Natural History and Science Bulletin. 41, 371-383.

Neotheropoda indet. (Rowe, 1989)
Sinemurian-Pliensbachian, Early Jurassic
Rock Head MNA 219-0, Silty Facies Member of the Kayenta Formation, Arizona, US
Material- (MNA.V.100; paratype of Syntarsus kayentakatae) partial ilium (Rowe, 1989)
(MNA.V.140; paratype of Syntarsus kayentakatae) humerus, femur, partial tibia, fragments (Rowe, 1989)
(MNA.V.3181) tooth fragment (?), two pubic fragments (Gay, 2001)
Comments- Rowe (1989) first mentions "a partial left ilium with the supra-acetabular crest (MNA V100), and weathered fragments of a femur and humerus ofa presumed juvenile (MNA V140)" which he refers to Syntarsus kayentakatae. Tykoski (1998) noted "The femur has a low, conical anterior trochanter" and "The bone texture is rough and pitted in the place a trochanteric shelf develops in robust adults", and concluded that while the locality matches kayentakatae the specimens "could just as conceivably be those of a very young Dilophosaurus wetherilli." He stated "they should be assigned to Ceratosauria indet.", which is equivalent to Neotheropoda here.
Gay (2001) described a specimen found in 1978, which he identified as "a distal humerus, a partial distal fibula and a tooth fragment" of an infant Dilophosaurus. He claimed "The humerus very closely resembles that of UCMP 37302, with the ectocondyle, ectepicondyle, entocondyle and entepicondyle all being present in the same arrangement ... but much more pronounced", the fibula has "the same general shape and curvature found in both MNA PI. 530 and MNA Pl.539", and "The tooth may or may not belong to this animal." However, Marsh and Rowe (2020) reidentified this specimen as "the distal end of a coelophysoid pubis", where the supposed humeral and fibular pieces are parts of a pubis (Marsh, pers. comm. 8-9-21) while the tooth may be incorrectly assigned.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9(2), 125-136.
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications for ceratosaurid phylogeny. Masters Thesis, University of Texas at Austin. 217 pp.
Gay, 2001. New specimens of Dilophosaurus wetherilli (Dinosauria: Theropoda) from the Early Jurassic Kayenta Formation of northern Arizona. Mesa Southwest Museum Bulletin. 8, 19-23.
Marsh and Rowe, 2020. A comprehensive anatomical and phylogenetic evaluation of Dilophosaurus wetherilli (Dinosauria, Theropoda) with descriptions of new specimens from the Kayenta Formation of northern Arizona. Journal of Paleontology. 94(Memoir 78), 103 pp.

undescribed Neotheropoda (Small, 2009)
Late Norian, Late Triassic
'red siltstone member' of the Chinle Formation, Colorado, US
Material- (DMNH coll.) ?cranial elements, pelvic elements, femora, pedal elements
Comments- Small (2009) wrote in an abstract "coelophysoid dinosaurs [are] represented by femora, pelvic material, pes material, and possible cranial bones", while Martz and Small (2019) said they "will be described in a future publication." Given the recent interpretation of coelophysoids as a basal grade of neotheropods and similarity to e.g. Liliensternus, it is assigned to Neotheropoda here.
References- Small, 2009. A Late Triassic dinosauromorph assemblage from the Eagle Basin (Chinle Formation), Colorado, U.S.A.. Journal of Vertebrate Paleontology. 29(3), 182A.
Martz and Small, 2019. Non-dinosaurian dinosauromorphs from the Chinle Formation (Upper Triassic) of the Eagle Basin, northern Colorado: Dromomeron romeri (Lagerpetidae) and a new taxon, Kwanasaurus williamparkeri (Silesauridae). PeerJ. 7:e7551.

undescribed possible neotheropod (Hunt, Huber, Reid, Frost, Cotton and Cotton, 1997)
Rhaetian, Late Triassic
Redonda Formation of the Dockum Group, New Mexico, US
Material- (NMMNH? coll.) teeth, partial femur, phalanx
Comments- One of two specimens (along with NMMNH P-22494) mentioned in an abstract, Hunt et al. (1997) state "The second specimen is only partially excavated and occurs stratigraphically just below the Redonda "ledge" -the uppermost lacustrine shoreline calcarenite in eastern Quay County. This specimen includes at least one partial femur, a podial and teeth. Both specimens represent theropods more derived than Herrerasauridae with hip heights of about 1 m. Neither specimen is generically determinate and both may represent the same taxon."
Reference- Hunt, Huber, Reid, Frost, Cotton and Cotton, 1997. Theropod dinosaurs from the latest Triassic Redonda Formation of east-central New Mexico. New Mexico Geological Society Annual Spring Meeting. 56.

unnamed neotheropod (Griffin, 2019)
Late Norian, Late Triassic
NMMNH L-110, Bull Canyon Formation of the Dockum Group, New Mexico, US
Material- (NMMNH P-4563) (subadult) proximal left fibula (59.43 mm anteropost)
References- Griffin, 2019. Large neotheropods from the Upper Triassic of North America and the early evolution of large theropod body sizes. Journal of Paleontology. 93(5), 1010-1030.
Griffin and Nesbitt, 2020 (2019 online). Does the maximum body size of theropods increase across the Triassic-Jurassic boundary? Integrating ontogeny, phylogeny, and body size. The Anatomical Record. 303, 1158-1169.

unnamed Neotheropoda (Cope, 1887a)
Late Norian, Late Triassic
Arroyo Seco, Petrified Forest Member of the Chinle Formation, New Mexico, US
Material- ?(AMNH 2702; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis) mid caudal vertebra (51 mm) (Cope, 1887a)
?(AMNH 2703; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis) manual ungual (Cope, 1887b)
(AMNH 2706; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis; paratype of Eucoelophysis baldwini) pubis (228 mm) (Cope, 1887b)
?(AMNH 2707; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis) manual phalanx I-1 (43 mm) (Cope, 1887b)
(AMNH 2708; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis; not lectotype of Coelophysis bauri, contra Welles, 1984) ilium (~155 mm) (Cope, 1887b)
?(AMNH 2715; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis) dorsal centrum (42 mm) (Cope, 1887a)
?(AMNH 2716; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis) ischium(?) (Cope, 1887b)
(AMNH 2717; = AMNH 2702 before 1973; paralectotype of Coelurus bauri) third or fourth cervical centrum (53 mm) (Cope, 1887a)
(AMNH 2718; = AMNH 2702 before 1973; paralectotype of Coelurus bauri) distal ischium (Cope, 1887b)
?(AMNH 2719; = AMNH 2702 before 1973; paralectotype of Coelurus bauri) proximal ischium(?) (Cope, 1887b)
(AMNH 2720; = AMNH 2702 before 1973; paralectotype of Coelurus bauri) partial fourth or fifth cervical vertebra (Cope, 1887b)
(AMNH 2722; ; = AMNH 2702A before 1973; lectotype of Coelurus bauri) incomplete sacrum (20, 19, 19, 16, ? mm), proximal tibia (Cope, 1887a)
?(AMNH 2723; = AMNH 2702 before 1973; paralectotype of Coelurus bauri) dorsal centrum (30 mm) (Cope, 1887b)
(AMNH 2724; = AMNH 2702 before 1973; paralectotype of Coelurus bauri) proximal pubis (Cope, 1887b)
?(AMNH 2727; = AMNH 2703 before 1973; paralectotype of Tanystropheus willistoni) distal caudal centrum (Cope, 1887b)
?(AMNH 2728; = AMNH 2706 before 1973) distal metatarsal IV(?) (Huene, 1915)
?(AMNH 2729; = AMNH 2706 before 1973) dorsal transverse process (Huene, 1915)
?(AMNH 2730; = AMNH 2706 before 1973) distal metatarsal III (Huene, 1906)
?(AMNH 2731; = AMNH 2706 before 1973) proximal pubis (Huene, 1915)
?(AMNH 2732; = AMNH 2706 before 1973) five vertebral fragments (Padian, 1986)
?(AMNH 2734; = AMNH 2705 before 1973) incomplete mid caudal vertebra (Huene, 1915)
?(AMNH 2735; = AMNH 2705 before 1973) partial distal caudal centrum (Cope, 1887b)
?(AMNH 2736; = AMNH 2705 before 1973) partial cervical neural arch (Huene, 1915)
?(AMNH 2737; = AMNH 2705 before 1973) proximal humerus (Huene, 1915)
?(AMNH 2738; = AMNH 2705 before 1973) ischial fragments (Huene, 1915)
?(AMNH 2739; = AMNH 2706 before 1973) dorsal centrum (Huene, 1915)
?(AMNH 2740; = AMNH 2705 before 1973) distal metatarsal III (Huene, 1915)
?(AMNH 2742; = AMNH 2705 before 1973) bone fragments (Padian, 1986)
?(AMNH 2743; = AMNH 2705 before 1973) vertebra, six fragments (Padian, 1986)
?(AMNH 2744) partial proximal caudal vertebra (Huene, 1906)
?(AMNH 2745) proximal fibula (Huene, 1915)
?(AMNH 2746; = AMNH 2707 before 1973) five caudal vertebra fragments (Padian, 1986)
?(AMNH 2747; = AMNH 2707 before 1973) pedal fragments (Padian, 1986)
?(AMNH 2748; = AMNH 2707 before 1973) vertebral fragments (Padian, 1986)
?(AMNH 2749; = AMNH 2704 before 1973) dorsal centrum (Huene, 1915)
?(AMNH 2750; = AMNH 2704 before 1973) anterior sacrum (Huene, 1915)
?(AMNH 2751; = AMNH 2704 before 1973) posterior cervical vertebra (Huene, 1915)
?(AMNH 2752; = AMNH 2704 before 1973) partial anterior dorsal vertebra (Huene, 1915)
?(AMNH 2753; = AMNH 2704 before 1973) vertebral and bone fragments (Padian, 1986)
? fragments (Williston and Case, 1912)
? fragmentary elements (Colbert, 1989)
Late Norian, Late Triassic
locality 1, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-453) incomplete dorsal vertebra (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
locality 2, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-456) dorsal vertebra (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
locality 5, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-462) distal ?fibula (Sullivan, Lucas, Heckert and Hunt, 1996)
?(SMP VP-465) distal ?femur (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
locality 6, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-469) dorsal vertebra, proximal tibia, distal fibula (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
locality 8, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-476) fragments (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
locality 9, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-478) three dorsal vertebrae, caudal vertebra, proximal tibia, proximal metatarsal? (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
locality 12, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-487) dorsal centrum (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
locality 13, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(SMP VP-491) distal scapula (Sullivan, Lucas, Heckert and Hunt, 1996)
Late Norian, Late Triassic
Canjilon Quarry UCMP V2816, Petrified Forest Member of the Chinle Formation, New Mexico, US
Material- (UCMP 152645) ilial fragment, partial astragali, calcaneum, distal tarsal III, phalanx I-1, partial metatarsal II, four partial pedal phalanges, pedal ungual (Nesbitt and Stocker, 2008)
?(UCMP coll.; lost) mid dorsal vertebra (Long and Murry, 1995)
Arroyo Seco material- The AMNH material including the lectotype of Coelurus (= Tanystropheus, = Coelophysis) bauri was collected by Baldwin with the lectotypes of Coelurus (= Tanystropheus, = Coelophysis) longicollis (AMNH 2701), Tanystropheus (= Coelophysis) willistoni (AMNH 2726) and the holotype of Longosaurus longicollis (AMNH 2705) in February 1881 "at three localities, one described as "Gallina Canyon", the other two as "Arroyo Seco"" (Sullivan et al., 1996). The latter reference further states that "It is not certain from exactly which of the three localities BALDWIN collected a given syntype or original referred specimen; the precise association was lost long ago." Thus "Arroyo Seco" as used on this site refers to any of those three localities. After preliminary descriptions by Cope (1887a, b), Huene (1906, 1915) described and illustrated most material, and Padian (1986) provided the definitive historical review of the specimens. Padian also first articulated the issue that the Arroyo Seco elements were less diagnostic than associated skeletons like UCMP 129618 found in 1982 or the numerous Coelophysis Quarry (= Ghost Ranch, = Whitaker Quarry) skeletons found in 1947. Hunt and Lucas (1991) attempted to solve this by naming the Coelophysis Quarry specimens Rioarribasaurus colberti, but the ICZN (1996) ruled that a Coelophysis Quarry specimen (AMNH 7224) is the neotype of Coelophysis bauri, leaving the Arroyo Seco specimens as not definitely Coelophysis. Sullivan and Lucas (1999) erected a new species of supposed ceratosaur sensu lato from near Arroyo Seco, Eucoelophysis baldwini, and believed Baldwin's Arroyo Seco material may belong to it, but stated "most of these specimens are not diagnostic because they lack apomorphic characters that would permit unambiguous generic and specific assignment." While Sullivan and Lucas referred Baldwin's AMNH 2706 pubis to the taxon and Heckert et al. (2000) proposed the Snyder Quarry coelophysid was Eucoelophysis, Nesbitt et al. (2005, 2007) and Ezcurra (2006) later demonstrated Eucoelophysis was not a theropod but instead a silesaur and that these referred materials are coelophysoid-grade theropods. Nesbitt et al. (2007) reexamined the Arroyo Seco 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 coelophysoid-grade theropods 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 supposedly undiagnostic within Archosauria, femur AMNH 2704 is a silesaur (this site), distal femur AMNH 2721 is Dromomeron romeri (Nesbitt et al., 2009), and distal femur AMNH 2725 is a shuvosaurid (Nesbitt et al., 2007). The remaining material is listed here for convenience pending restudy.
Williston and Case (1912) reported "bone fragments referred provisionally to the genus Coelophysis" were found in the "immediate locality" of the type material. Colbert (1989) stated "In the summer of 1986, some fragmentary Coelophysis bones were found at Ghost Ranch, somewhat to the east of Arroyo Seco and downstream from the Ghost Ranch Quarry. This locality is about a mile to the southeast of the quarry." Sullivan (1994) reported "indeterminate ceratosaur fossils which are considered topotypic material of Coelophysis bauri" found in 1993 from five sites around Arroyo Seco. Sullivan et al. (1996) published the specimens and localities, again proposing them as topotypes for Coelophysis bauri. These are undescribed with only SMP VP-487 figured, and are probably indeterminate at levels between Coelophysidae and Archosauromorpha.
Canjilon material- Long and Murry (1995) stated "A complete dorsal vertebra (UCMP V2816) was found among the numerous Typothorax and pseudopalatine phytosaur remains at the Canjilon Quarry", and "Though this vertebra is comparable to those of AMNH 7224 in size and detail, we believe this slightly older theropod specimen cannot be determined to genus and refer it to Theropoda incertae sedis." Note V2816 is the locality number for Canjilon Quarry, not a specimen number. Based on this reported resemblence to Coelophysis, this is placed in Neotheropoda here instead of Saurischia because herrerasaurids and Chindesaurus have much shorter dorsal centra although they were classified as theropods by Long and Murry. Angielczyk (2002) reports "This specimen could not be found."
Discovered on October 13 1928, UCMP 152645 was first published by Nesbitt and Stocker (2008) who briefly describe and figure the unfused proximal tarsus. They conclude "The combination of the character states (perforated acetabulum, box-like calcaneum, and anteroposteriorly compressed ascending process of the astragalus) confirm that UCMP 152645 represents a theropod." Within that clade, it seems to be basal as "The ascending process has an anteroposterior thickness more similar to those of Coelophysis bauri (AMNH FR 30576), the 'Padian theropod' (UCMP 129618; Padian, 1986) and GR 211 [the Hayden Quarry coelophysid], than to the much more anteroposteriorly thin process of Dilophosaurus."
References- Cope, 1887a. The dinosaurian genus Coelurus. American Naturalist. 21, 367-369.
Cope, 1887b. A contribution to the history of the Vertebrata of the Trias of North America. Proceedings of the American Philosophical Society. 24(126), 209-228.
Huene, 1906. Ueber die Dinosaurier der Aussereuropaischen Trias. Geologische und Paläontologische Abhandlungen. 12, 99-156.
Williston and Case, 1912. The Permo-Carboniferous of northern New Mexico. The Journal of Geology. 20(1), 1-12.
Huene, 1915. On reptiles of the New Mexican Trias in the Cope collection. Bulletin American Museum of Natural History. 34, 485-507.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
Padian, 1986. On the type material of Coelophysis Cope (Saurischia: Theropoda) and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation). In Padian (ed.). The Beginning of the Age of Dinosaurs: Faunal Change Across the Triassic-Jurassic Boundary. Cambridge University Press. 45-60.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic dinosaur from New Mexico (USA). Paläontologische Zeitschrift. 65(1/2), 191-198.
Sullivan, 1994. Topotypic material of Coelophysis bauri (Cope) and the Coelophysis-Rioarribasaurus-Syntarsus problem. Journal of Vertebrate Paleontology. 14(3), 48A.
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.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the southwestern Unites States. New Mexico Museum of Natural History and Science Bulletin. 4, 1-254.
Sullivan, Lucas, Heckert and Hunt, 1996. The type locality of Coelophysis, a Late Triassic dinosaur from north-central New Mexico (USA). Paläontologische 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.
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.
Angielczyk, 2002. A selective annotation of published Triassic vertebrates from the UCMP collection. In Heckert and Lucas (eds.). Triassic Stratigraphy and Paleontology. Bulletin of the New Mexico Museum of Natural History and Science. 21, 297-301.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. Journal of Vertebrate Paleontology. 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 and Stocker, 2008. The vertebrate assemblage of the Late Triassic Canjilon Quarry (northern New Mexico, USA), and the importance of apomorphy-based assemblage comparisons. Journal of Vertebrate Paleontology. 28(4), 1063-1072.
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.

unnamed Neotheropoda (Chatterjee, 1993)
Middle Norian, Late Triassic
Post Quarry MOTT 3624, Lower Cooper Canyon Formation of the Dockum Group, Texas, US
Material- (TTU-P10071) incomplete ilium (Lehane, 2005)
(TTU-P11044) tibia (111 mm) (Chatterjee, 1993)
Middle Norian, Late Triassic
Davidson Creek UCMP V6333, Lower Cooper Canyon Formation of the Dockum Group, Texas, US
?(UCMP 65737; field number 1963/2) cervical vertebral fragments (UCMP online)
?(UCMP 65773; field number 1963/22) dorsal vertebra (UCMP online)
?(UCMP 65774; field number 1963/22) caudal vertebra (UCMP online)
?(UCMP 65775; field number 1963/22) (UCMP online)
Comments- Chatterjee (1993) stated as correspondence at the end of his Shuvosaurus description "we have just found a diagnostic leg bone (tibia) from the Post Quarry which confirms the presence of an ostrich dinosaur", though this was not technically referred to the genus. The tibia was later assigned the number TTU-P11044, as revealed in Lehane's (2005) thesis, who suggested it was too small to belong to Shuvosaurus individuals known from that quarry. He assigned it to Theropoda indet., as did Nesbitt and Chatterjee (2008) who described it as Theropoda indet., and said it is "identical to the Snyder Quarry coelophysoid (NMMNH P-29046, NMMNH P-29047, and NMMNH P-31293; Nesbitt et al. 2007) and Coelophysis bauri (AMNH 7223; AMNH 7224) tibiae." Marsh et al. (2019) suggested it may belong to the Chindesaurus + Tawa "clade owing to the presence of two notches on the posterior margin of the proximal end", but the second concavity is far medial of Chindesaurus or Tawa, and the tibia otherwise differs in the long cnemial crest, fibular crest, anteriorly shifted lateral condyle, and concave posterolateral edge in distal view, which are all more similar to neotheropods (Martz et al., 2012). Contra Nesbitt and Chatterjee, it is not identical to the Snyder Quarry theropod or Coelophysis bauri, being most similar to Upper Cooper Canyon tibia TTU-P10534 in having an anteroposteriorly deep distal end (97% of width), short posterolateral process, anteriorly placed lateral condyle and no medial malleolus. It differs from that tibia in being even deeper anteroposteriorly and having a convex anteromedial edge (both in distal view).
Lehane (2005) referred incomplete ilium TTU-P10071 to Shuvosaurus in his thesis, but it was identified as Coelophysis by Lehman and Chatterjee (2005) and agreed to be theropod by later authors (Nesbitt and Chatterjee, 2008; Martz et al., 2012). While Martz et al. only identify it to the level of Neotheropoda, it is probably a coelophysid based on the striaght dorsal edge.
UCMP 65737 was collected on June 15 1963, while UCMP 65773-65775 were collected on June 22 1963. These were all referred to ?Coelophysis on the UCMP online catalogue, but could easily be from e.g. shuvosaurids pending further information.
References- Chatterjee, 1993. Shuvosaurus, a new theropod. National Geographic Research and Exploration. 9(3), 274-285.
Lehane, 2005. Anatomy and relationships of Shuvosaurus, a basal theropod from the Triassic of Texas. Masters thesis, Texas Tech University. 92 pp.
Lehman and Chatterjee, 2005. Depositional setting and vertebrate biostratigraphy of the Triassic Dockum Group of Texas. Journal of Earth System Science. 114(3), 325-351.
Nesbitt and Chatterjee, 2008. Late Triassic dinosauriforms from the Post Quarry and surrounding areas, west Texas, U.S.A. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen. 249(2), 143-156.
Martz, Mueller, Nesbitt, Stocker, Parker, Atanassov, Fraser, Weinbaum and Lehane, 2012. A taxonomic and biostratigraphic re-evaluation of the Post Quarry vertebrate assemblage from the Cooper Canyon Formation (Dockum Group, Upper Triassic) of southern Garza County, western Texas. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103, 1-26.
Marsh, Parker, Langer and Nesbitt, 2019. Redescription of the holotype specimen of Chindesaurus bryansmalli Long and Murry, 1995 (Dinosauria, Theropoda), from Petrified Forest National Park, Arizona. Journal of Vertebrate Paleontology. 39(3), e1645682.

unnamed neotheropod (Sarigül, 2014)
Late Norian, Late Triassic
Headquarters South MOTT 3898, Middle Cooper Canyon Formation of the Dockum Group, Texas, US
Material- (TTU-P14786) distal tibia
Comments- Discovered in the 2000s, Sarigül (2017) wrote "a deeply penetrated articulation facet on the rhomboidal distal surface for the ascending process of the astragalus, an enlarged posterior process on the lateral side, and a characteristic longitudinal ridge on the posterior side strongly support the neotheropod affinity." Indeed, it is very similar to other Cooper Canyon tibiae (TTU-P10071, P10534) in having an anteroposteriorly deep distal end (90% of width), short posterolateral process, and almost no medial malleolus.
References- Sarigül, 2014. Anatomy of the Late Triassic dinosauromorphs from the Dockum Group of Texas: Their biostratigraphic, paleobiogeographic and evolutionary significance. PhD thesis. Texas Tech University. 300 pp.
Sarigül, 2017. New theropod fossils from the Upper Triassic Dockum Group of Texas, USA, and a brief overview of the Dockum theropod diversity. PaleoBios. 34, 1-18.

unnamed neotheropod (Cunningham, Hungerbuhler, Chatterjee and McQuilkin, 2002)
Rhaetian, Late Triassic
Patty East (Patricia Site) MOTT 3880, Upper Cooper Canyon Formation of the Dockum Group, Texas, US
Material- (TTU-P10534) tibia (~253 mm)
Comments- This was discovered between 2001 and 2002. Cunningham et al. (2002) identified TTU-P10534 as "An isolated tibia is referred to a large ornithischian dinosaur", 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 basal neotheropods. While it is listed as unnumbered in Nesbitt et al.'s paper, the number is listed in Nesbitt and Chatterjee (2008). It was assigned to Neotheropoda by Sarigul (2017) and stated to closely resemble Dilophosaurus in "the robust and poorly flared cnemial crest, slightly elevated medial border of the proximal surface, relative sizes of tibial condyles, the triangular fibular crest and the shape of the distal articular surface." On the other hand, Nesbitt et al. claimed "it is indistinguishable from the tibia of Gojirasaurus and the smaller, less robust, tibiae of Coelophysis." However, it differs from named basal neotheropods in it anteroposteriorly deep distal end (87% of width) and short posterolateral process, while the anteriorly placed lateral condyle is unlike Gojirasaurus and the Snyder Quarry theropod, but like Dilophosaurus and the Petrified Forest theropod. It may be close to Lepidus in the lack of a medial malleolus, as that taxon also seems to have a short posterolateral process but is unavailable in distal view.
References- Cunningham, Hungerbuhler, Chatterjee and McQuilkin, 2002. Late Triassic vertebrates from the Patricia Site near Post, Texas. Journal of Vertebrate Paleontology. 22(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.
Nesbitt and Chatterjee, 2008. Late Triassic dinosauriforms from the Post Quarry and surrounding areas, west Texas, U.S.A. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen. 249(2), 143-156.
Sarigül, 2014. Anatomy of the Late Triassic dinosauromorphs from the Dockum Group of Texas: Their biostratigraphic, paleobiogeographic and evolutionary significance. PhD thesis. Texas Tech University. 300 pp.
Sarigül, 2017. New theropod fossils from the Upper Triassic Dockum Group of Texas, USA, and a brief overview of the Dockum theropod diversity. PaleoBios. 34, 1-18.

undescribed neotheropod (Galton, 1976)
Early Hettangian, Early Jurassic
Shuttle Meadow Formation, Connecticut, US
Material- tooth
Comments- Galton (1976) stated "A small tooth referable to Coelophysis was collected in 1970 by B. Cornet and N. G. McDonald (personal communication) from black shale in a stream cut in the Shuttle Meadow Formation, northeast side of Totoket Mountain, North Guilford, Connecticut (see Cornet et al., 1973, locality l )." However, Coelophysis teeth are not known to be diagnostic and dental characters have not been suggested to distinguish taxa of basal neotheropods..
Reference- Galton, 1976. Prosauropod dinosaurs (Reptilia: Saurischia) of North America. Postilla. 169, 1-98.

unnamed neotheropod (Simms, Smyth, Martill, Collins and Byrne, 2020 online)
Early Hettangian?, Early Jurassic
Planorbis Zone(?) of the Waterloo Mudstone Formation(?), Lias Group, Northern Ireland
Material- (BELUM K12493) tibial fragment
Comments- This was found on April 15 1981 and described by Simms et al. (2020 online), who considered it most similar to the coeval Sarcosaurus.
Reference- Simms, Smyth, Martill, Collins and Byrne, 2020 online. First dinosaur remains from Ireland. Proceedings of the Geologists' Association. Article in Press. DOI: 10.1016/j.pgeola.2020.06.005

unnamed possible neotheropod (Rioult, 1978)
Early Hettangian, Early Jurassic
Calcaire de Valognes, Manche, France
?(University of Caen coll.; destroyed) tooth
Comments- Buffetaut et al. (1991) mentions "A tooth referred to Megalosaurus cloacinus Quenstedt, from the Lower Hettangian of the Calcaire de Valognes at Valognes (Manche), [which] has been mentioned by Rioult (1978a) as having been destroyed by an air raid on the University of Caen in 1944." Without additional details, it can only be said that the timing suggests a neotheropod.
References- Rioult, 1978. Inventaire des dinosauriens mésozoïques de Normandie. Ecosystèmes continentaux mésozoiques de Normandie (Livret-guide). Université de Caen. 26-29.
Buffetaut, Cuny and Le Loeuff, 1991. French Dinosaurs: The best record in Europe? Modern Geology. 16(1/2), 17-42.

unnamed Neotheropoda (Delsate, 2000)
Late Hettangian, Early Jurassic
Feidt Quarry, Luxembourg Sandstone Formation, Luxembourg
Material- (MHNL BR778) incomplete pedal phalanx III-1 (Delsate, 2000)
(MHNL BR924) lateral tooth (Delsate and Ezcurra, 2014)
Comments- MHNL BR778 was first identified as a manual phalanx III-3 by Delsate (2000) before being redescribed by Delsate and Ezcurra (2014) as a pedal phalanx.
References- Delsate, 2000. Paléontologie des vertébrés au Grand-Duché de Luxembourg: Découvertes récentes et travaux en cours. Archives de l’Institut Grand-Ducal de Luxembourg, Section des Sciences Naturelles, Physiques et Mathématiques, Nouvelle Série. 43, 49-54.
Delsate and Ezcurra, 2014. The first Early Jurassic (Late Hettangian) theropod dinosaur remains from the Grand Duchy of Luxembourg. Geologica Belgica. 17(2), 175-181.

unnamed possible neotheropod (Huene, 1921)
Middle Norian, Late Triassic
Middle Löwenstein Formation, Germany
Material- (SMNS 12670 in part) quadrate, pterygoid, cervical vertebra, mid dorsal vertebra, scapula, incomplete ilium, pubis
Comments- Huene (1921) referred these to Aetosaurus, but they were reidentified by Walker (1961) as a 'coelurosaur' similar to Procompsognathus. The quadrate was originally identified as a pubis, the pterygoid as a postorbital+postfrontal, the mid dorsal as a proximal caudal, the scapula as an ischium, and the pubis as a scapula. Knoll (2008) considered most of the material could be aetosaurian or indeterminate, but thought the scapula might be theropodan though not Procompsognathus. Huene also describes the pubis SMNS 12596 as Saltoposuchus, which Walker (1970) referred to a 'coelurosaur' resembling Procompsognathus. This was figured as a crocodylomorph by Knoll (2008).
References- Huene, 1921. Neue Pseudosuchier und Coelurosaurier aus dem württembergischen Keuper. Acta Zoologica. 2, 329-403.
Walker, 1961. Triassic reptiles from the Elgin area: Stagonolepis, Dasygnathus and their allies. Philosophical Transactions of the Royal Society of London, Series B. 244, 103-204.
Walker, 1970. A revision of the Jurassic reptile Hallopus victor (Marsh), with remarks on the classification of crocodiles. Philosophical Transactions of the Royal Society of London, Series B. 257, 323-372.
Knoll, 2008. On the Procompsognathus postcranium (Late Triassic, Germany). Geobios. 41(6), 779-786.

undescribed averostran (Dong, Zhou and Zhang, 1983)
Toarcian?, Early Jurassic?
Shejiaju coal mine, Daanzhai Member of Ziliujing Formation, Sichuan, China
Material- (moderate to large) vertebrae, scapula
Comments- When discussing the Daanzhai Member, Dong et al. (1983) write "In 1978, Yihong Zhang collected a series of large vertebrae and a scapulae from the upper limestones at the Shejiaju coal mine, Rongjing Co. Preliminary analysis diagnoses the specimens as a moderate-sized carnosaur" (translated). Rongjing County is now known as Yingjing County, and Shejiaju does not refer to the city in Hunan or the village in Hubei. The material has never been described. Considering the size and 'carnosaur' label, it may be a basal averostran like Sinosaurus and Saltriovenator or just outside the clade like Cryolophosaurus and Sarcosaurus.
Reference- Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica Sinica. Whole Number 162, New Series C, 23, 136 pp.

undescribed neotheropod (Liston, Naish, Hone, Tianyang and Jian-Rong, 2014)
Hettangian, Early Jurassic
DaWaShan, Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
Material- five partial teeth
Comments- Liston et al. (2014) state these are distinct from Sinosaurus and "all other theropod taxa currently known from the Lufeng fauna."
Reference- Liston, Naish, Hone, Tianyang and Jian-Rong, 2014. New data on Early Jurassic theropod diversity and feeding behavior in the Lufeng Formation of Yunnan, China. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 169.

unnamed possible neotheropod (Simmons, 1965)
Sinemurian, Early Jurassic
Ta Ti, Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Material- (FMNH CUP 2091) distal femur
Comments- Simmons (1965) identified FMNH CUP 2091 as podokesaurid, but it lacks an infrapopliteal ridge unlike coelophysoids, and differs from most basal theropods in several characters (concave medial margin in distal view; concave posterior margin of medial condyle in distal view; ectocondylar tuber projects far posterior to medial condyle and is angled medially).
References- Young, 1951. The Lufeng saurischian fauna in China. Palaeontologica Sinica. C(13), 1-96.

unnamed neotheropod (Novas, Chatterjee, Ezcurra and Kutty, 2009; described by Novas, Ezcurra, Chatterjee and Kutty, 2010)
Rhaetian, Late Triassic
Lower Dharmaram Formation, India
Material- (ISI R283) incomplete femur
Comments- This was initially called a coelophysoid (Novas et al., 2009) before being described as a non-averostran neotheropod by Novas et al. (2010).
References- Novas, Chatterjee, Ezcurra and Kutty, 2009. New dinosaur remains from the Late Triassic of Central India. Journal of Vertebrate Paleontology. 29(3), 156A.
Novas, Ezcurra, Chatterjee and Kutty, 2010. New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of central India. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 333-349.

undescribed Neotheropoda (Allain, Aquesbi, Dejax, Meyer, Monbaron, Montenat, Richir, Rochdy, Russell and Taquet, 2004)
Toarcian, Early Jurassic
O locality, Douar of Tazouda, Upper bone-bed of the Toundoute continental series, Morocco
Material- (MHNM-O coll.) (Peyer and Allain, 2010)
Toarcian, Early Jurassic
R locality, Douar of Tazouda, Upper bone-bed of the Toundoute continental series, Morocco
(MHNM-R coll.) (Peyer and Allain, 2010)
Toarcian, Early Jurassic
To2 locality, Douar of Tazouda, Lower bone-bed of the Toundoute continental series, Morocco
(MHNM-To2 coll.) (large) phalanges (Allain, Aquesbi, Dejax, Meyer, Monbaron, Montenat, Richir, Rochdy, Russell and Taquet, 2004)
Comments- Discovered in 2000, Allain et al. (2004) noted that associated with the Tazhoudasaurus types "were isolated elements of medium-sized and large theropods of uncertain affinities", the former eventually being described as Berberosaurus and the latter yet undescribed. Allain et al. (2007) mention this as "a large carnivorous dinosaur of uncertain affinities." Peyer and Allain (2010) refer to this specimen when they mention "the lower bone bed has only yielded dinosaur remains at the To2 site, including one juvenile and a subadult skeleton of Tazoudasaurus and an enigmatic theropod." Pameiro (pers. comm. 7-2023) indicates by 2010 only phalanges of the large theropod were known.
Peyer and Allain also say "Current fieldwork at the new sites O and R promise further remains of Tazoudasaurus skeletons and theropod remains, all of which are still being excavated and prepared."
References- Allain, Aquesbi, Dejax, Meyer, Monbaron, Montenat, Richir, Rochdy, Russell and Taquet, 2004. A basal sauropod dinosaur from the Early Jurassic of Morocco. Comptes Rendus Palevol. 3, 199-208.
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.
Peyer and Allain, 2010. A reconstruction of Tazoudasaurus naimi (Dinosauria, Sauropoda) from the late Early Jurassic of Morocco. Historical Biology. 22(1-3), 134-141.

undescribed neotheropod (Choiniere, Benson, Botha, Barrett, Bordy, Chapelle, Dollman, Suarez, Viglietti, Sciscio and Butler, 2020)
Middle Norian-Rhaetian, Late Triassic
Lower Elliot Formation, South Africa
Material- incomplete cervical vertebra (~105 mm), partial cervical vertebra
Comments- These are highly elongate with anterior and posterior pleurocoels.
Reference- Choiniere, Benson, Botha, Barrett, Bordy, Chapelle, Dollman, Suarez, Viglietti, Sciscio and Butler, 2020. Taxonomically rich Late Triassic faunas from South Africa's lowermost Elliot Formation. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 105-106.

unnamed neotheropod (Munyikwa and Raath, 1999)
Pliensbachian, Early Jurassic
Paradys Farm, Upper Elliot Formation, South Africa
Material- (BP/1/5278) (juvenile) premaxillae, incomplete maxillae, partial nasals, partial dentaries, teeth
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.
Comments- This was discovered in 1985. Initially described as a specimen of Syntarsus rhodesiensis by Munyikwa and Raath (1999), and tentatively referred to Dracoventor by Yates (2005) in his description of that taxon. Ezcurra (2012) found this to be a non-coelophysid coelophysoid (in a sense including Liliensternus but not Dilophosaurus) in a large unpublished analysis, while Dracovenator was still a dilophosaurid. Wang et al. (2017) found this in a polytomy with Camposaurus, Megapnosaurus and kayentakatae within Coelophysidae.
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.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.

undescribed neotheropod (Blackbeard and Yates, 2007)
Pliensbachian, Early Jurassic
Spioenkop Farm, Upper Elliot Formation, South Africa
Material- (BPI/1/coll.) (large) teeth
Reference- Blackbeard and Yates, 2007. The taphonomy of an Early Jurassic dinosaur bonebed in the Northern Free State (South Africa). Journal of Vertebrate Paleontology. 27(3), 49A.

undescribed Neotheropoda (Slaughter, Hickerson and Hammer, 1994)
Rhaetian-Toarcian, Late Triassic-Early Jurassic
Hanson Formation, Antarctica
Material- (FMNH coll.) (multiple taxa) teeth
Comments- Hammer et al. (1994) write "A recent review (Slaughter, Hickerson, and Hammer 1994) of serration densities and patterns of teeth found near gnawed elements among the Falla Formation fossils showed that the teeth represented at least two (and possibly three) different types of scavenging theropods." Although described as 'halticosaurid' by Ford (DML, 1998), coelophysoid teeth are not known to be diagnostic.
References- Hammer, Hickerson and Slaughter, 1994. A dinosaur assemblage from the Transantarctic Mountains. Antarctic Journal. 29(5), 31-33.
Slaughter, Hickerson and Hammer, 1994. Analysis of Antarctic theropod teeth based on serration densities and patterns. Geological Society of America Abstracts with Programs. 26, 61.
Ford, DML 1998. https://web.archive.org/web/20191030050824/http://dml.cmnh.org/1998Aug/msg00810.html

Coelophysis? willistoni (Cope, 1887) Cope, 1889
= Tanystophaeus willistoni Cope, 1887
Late Norian, Late Triassic
Arroyo Seco, Petrified Forest Member of the Chinle Formation, New Mexico, US
Lectotype- (AMNH 2726; = AMNH 2703 before 1973) partial ilium
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 Coelophysoidea.
References- Cope, 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proceedings of the American Philosophical Society. 24(126), 209-228.
Cope, 1889. On a new genus of Triassic Dinosauria. The American Naturalist. 23, 626.

"Comanchesaurus" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
"C. kuesi" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
Late Norian, Late Triassic
NMMNH L-110, Bull Canyon Formation of the Dockum Group, New Mexico, US
Material- (NMMNH P4569; Bull Canyon neotheropod) (subadult) incomplete atlantal intercentrum, partial anterior cervical epipophysis, two cervical central fragments, cervical rib tuberculum, dorsal centrum, two dorsal prezygapophyses, two dorsal transverse processes, four dorsal rib tubercula, five dorsal rib capitula, partial proximal caudal centrum, partial distal caudal centrum, fourteen partial dorsal or caudal centra, many dorsal or caudal central fragments, apical dorsal/sacral/caudal neural spine, left scapular fragment, two distal right manual phalanges, proximal manual ungual, right ilial pubic peduncle, right ilial supracetabular crest, proximal right ischial fragment, proximal left femur (65.41 mm wide trans), distal left tibia, distal left fibula, partial left astragalocalcaneum, distal right metatarsal II, distal left metatarsal IV, incomplete pedal phalanx II/III-1, proximal phalanx II/III-1, phalanx IV-4, three proximal ?pedal phalanges, distal pedal phalanx
Comments- NMMNH P4569 was originally seen as a coelophysoid by Lucas et al. (1985) and Hunt and Lucas (1989), though Murry and Long (1989) and Long and Murry (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. Griffin (2019) described the specimen in detail as "Neotheropoda genus indeterminate species indeterminate", placing it in the clade due to astraglocalcanear fusion and the presence of a ridge on the proximomedial fibula.
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 east-central 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. in Lucas and Hunt (eds.). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. 72-101.
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. in Lucas and Hunt (eds.). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. 29-64.
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). PhD thesis, University of New Mexico. 403 pp.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the southwestern United States. New Mexico Museum of Natural History and Science Bulletin. 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.
Griffin, 2019. Large neotheropods from the Upper Triassic of North America and the early evolution of large theropod body sizes. Journal of Paleontology. 93(5), 1010-1030.
Griffin and Nesbitt, 2020 (2019 online). Does the maximum body size of theropods increase across the Triassic-Jurassic boundary? Integrating ontogeny, phylogeny, and body size. The Anatomical Record. 303, 1158-1169.

Dolichosuchus Huene, 1932
D. cristatus Huene, 1932
Early Norian, Late Triassic
Kaltental, Löwenstein Formation, Germany
Holotype- (NHMUK 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. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.

Dracoraptor Martill, Vidovic, Howells and Nudds, 2016
D. hanigani Martill, Vidovic, Howells and Nudds, 2016
Early Hettangian, Early Jurassic
Blue Lias Formation, Wales
Holotype- (NGM 2015.5G1-11) (~2 m juvenile) partial skull, fragmentary mandibles, six teeth, ?hyoid, two cervical vertebrae, several incomplete cervical ribs, dorsal centrum, two dorsal ribs, gastralia, partial last sacral vertebra, partial first caudal vertebra, incomplete second caudal centrum, partial third caudal vertebra, partial fourth caudal vertebra, proximal caudal neural arch, mid caudal centrum, two proximal chevrons, distal humerus, radius (71 mm), ulna (73 mm), metacarpal II (~34 mm), phalanx II-1, phalanx III-?, manual ungual, phalanx IV-?, incomplete pubes (~212 mm), ischium (~150 mm), incomplete femur, tibial fragment, incomplete fibula, proximal metatarsal II, phalanx II-?, phalanx III-1, proximal metatarsal IV, two pedal unguals, two phalanges
....(NGM 2015.10G.1) two dorsal centra, neural arch fragment, partial astragalus, calcaneum?, distal tarsal III, distal tarsal IV, distal metatarsal I, metatarsal II (102 mm), proximal phalanx II-1, metatarsals III (116 mm), proximal phalanx III-1, metatarsal IV (93 mm), phalanx IV-1, four phalanges (two proximal), pedal ungual, metatarsal V
Diagnosis- (proposed) posteroventral medial premaxillary process (also in coelophysids); narial fossa absent on premaxilla; portion of maxilla anterior to antorbital fenestra longer than portion under fenestra (also in Eodromeus); maxilla articulates in slot on lateral jugal.
Other diagnoses- Martill et al. (2016) listed "large external naris with slender subnarial bar" in their diagnosis, but as the nasal is not definitely identified and has no narial border, the naris is of uncertain size. Furthermore, the slender subnarial process of the premaxilla is found in most basal saurischians (except Herrerasaurus, Daemonosaurus and Chilesaurus). Three premaxillary teeth are unique among the coelophysoids that Martill et al. classified Dracoraptor as, but shared with Daemonosaurus and Chilesaurus. The jugal process is no more slender than coelophysids, and not that different from Daemonosaurus. Almost all basal saurischians have an anteriorly directed pubis (except some herrerasaurids and Chilesaurus), and Dracoraptor's ischium is actually longer compared to its pubis than coelophysids (~71% vs. 61-~67%) and comparable to e.g. Herrerasaurus and Dilophosaurus. Finally, Martill et al. list "large dorsal process on distal tarsal IV" but there is no obvious dorsal process. The tarsal is very close to rhodesiensis in shape except the posterior third is narrower and it has a slightly concave medial edge.
Comments- This specimen was discovered in 2014 and was described in 2016.
Note the maxillae are in medial view, as the figure caption states, but contra the text. Contra the text and reconstruction, only the small anteriormost jugal groove probably articulated with the maxilla. The longer ridge and groove on the main body is common in early dinosaurs (e.g. Herrerasaurus) and non-articulating. The supraoccipital is in anterior (internal) view, not ventral view as stated by the text. Note the large posttemporal fenestrae as in Silesaurus but unlike dinosaurs. The cervical is not opisthocoelous, contra the text, as the anterior central surface is slightly concave. The supposed first caudal is near certainly a ?last sacral based on the broad transverse processes originating on the centrum (compare to e.g. Staurikosaurus). The authors do call it "a partially sacralised element", but any ambiguity seems unnecessary. The next element could easily be a sacral too, though its more fragmented condition makes this more uncertain. I'm doubtful the supposed furcula is correctly identified. One side is much narrower than the other, and each is curved in a different direction (thin side concave toward the outside of the angle). Furcular arms are subequal in width, and those of coelophysoids (e.g. kayentakatae) are basically circular in section, so that twisting in Dracoraptor is not an excuse. It's more probably a posterior dorsal rib, which are also similar in having a ridge along the outside corner. The tuberculum may be covered in matrix. The authors say "A calcaneum is not present. Two distal tarsals (dt III & dt IV) and part of a putative third are present in a row." No archosauriforms actually have three distal tarsals per pes (certainly no theropods do), so that 'putative third' is more likely the supposedly missing calcaneum, especially as it's placed right next to distal tarsal IV (labeled 'Ldtii'). The supposed "?Metaacarpal of digit I" [sic] is a metacarpal II, very similar to rhodesiensis, more elongate than metacarpal I and more robust than metacarpal III. While I haven't identified all of the phalanges in this block, it's clear Martill et al.'s statement "they are assumed to be from the left manus as they are associated with the left radius and ulna" is in error. For instance, the phalanx underlying the proximal radius is too large to belong to any manual digit and is probably pedal phalanx III-1, while supposed manual unguals I and III lack flexor tubercles ("I" shows an obvious depression in that area) and at least "III" is virtually straight. These unguals more nearly match pedal unguals of e.g. Coelophysis and Liliensternus, while supposed ungual II is manual due to its curvature and large flexor tubercle. Among other phalanges, that at the distal end of metacarpal II matches a manual phalanx II-1, that on the proximal end of metacarpal II belongs to manual digit III, that between unguals "I" and "II" looks like its from pedal digit II, and the small one by the anterior end of the dorsal centrum would be manual IV-1. The latter suggests a less reduced digit IV than coelophysids or Herrerasaurus. Thus the manual reconstruction with its short penultimate phalanges and metacarpal ratios should not be trusted.
Martill et al. (2016) recovered Dracoraptor as a basal coelophysoid sister to Coelophysidae using a version of Nesbitt et al.'s basal dinosaur matrix. 15% of their Dracoraptor entries are miscoded though, and once corrected and placed in an improved version of that matrix that excludes the highly modified Velociraptor OTU, it emerges as sister to a Daemonosaurus+Chilesaurus clade, which is itself sister to Neotheropoda.
References- Howells, Nudds, Martill, Vidovic, Hannigan and Hannigan, 2015. A new Early Jurassic theropod from Wales: Geological and discovery context. SVPCA 2015 abstracts, 43.
Martill, Vidovic, Howells and Nudds, 2015. The oldest Jurassic dinosaur: A basal neotheropod from the Hettangian of Great Britain. SVPCA 2015 abstracts, 52.
Martill, Vidovic, Howells and Nudds, 2016. The oldest Jurassic dinosaur: A basal neotheropod from the Hettangian of Great Britain. PLoS ONE. 11(1), e0145713.

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
Late Norian, Late Triassic
Revuelto Creek UCM 82021, Bull Canyon Formation of the Dockum Group, New Mexico, US
Holotype- (UCM 47221; in part) (5.5 m) pubis (497 mm), tibia (469 mm), metatarsal V (105 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. Griffin (2019) described the previously unrecognized metatarsal V.
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 demonstrated that Shuvosaurus is a pseudosuchian 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 Coelophysis rhodesiensis, "M." kayentakatae and Segisaurus, more derived than Coelophysis bauri. 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. Ezcurra and Novas (2007) found it to be the sister taxon of "Syntarsus" 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. Ezurra (2012) found Gojirasaurus to be a non-coelophysid coelophysoid sensu stricto in a large unpublished analysis.
References- Parrish and Carpenter, 1986. A new vertebrate fauna from the Dockum Formation (Late Triassic) of eastern New Mexico. In Padian (ed.). The Beginning of the Age of Dinosaurs. Cambridge University Press. 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 of Natural History and Science Bulletin. 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.
Gay, 2000 DML. https://web.archive.org/web/20191030050834/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 (eds.). The Dinosauria Second Edition. University of California Press. 47-70.
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. Journal of Vertebrate Paleontology. 25(3), 96A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Thesis. 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.
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.
Ezcurra and Novas, 2007 (online 2006). Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
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.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Griffin, 2019. Large neotheropods from the Upper Triassic of North America and the early evolution of large theropod body sizes. Journal of Paleontology. 93(5), 1010-1030.

Halticosauridae Bock, 1952
Halticosaurinae Bock, 1952 vide 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 and/or averostrans/tetanurines. 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. 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. 464 pp.
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, 1908
H. longotarsus Huene, 1908
Middle Norian, Late Triassic
Pfaffenhofen, Middle Löwenstein Formation, Germany
Holotype- (SMNS 12353) incomplete dentary (lost), partial anterior cervical vertebra (lost), incomplete ?sixth cervical vertebra (53 mm), two fragmentary cervical vertebrae (lost), dorsal centrum (43 mm), partial dorsal centrum, incomplete fused second (35 mm) and third (33 mm) sacral vertebrae (lost), proximal caudal vertebra (36 mm), proximal humerus (lost), ilial fragment (lost), proximal femora (~223 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- The holotype was discovered in 1906. 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. Geologische und Palaeontologische Abhandlungen Suppl. 1(1), 1-419.
Huene, 1909. Skizze zu einer Systematik und Stammesgeschichte der Dinosaurier. Centralblatt für Mineralogie, Geologie und Paläontologie. 1909, 12-22.
Zittel, 1911. Grundzüge der Paläontologie (Paläozoologie). II. Abteilung. Vertebrata. Druck und Verlag von R. Oldenbourg, München. 1-598.
Huene, 1914. Das natürliche System der Saurischia. Centralblatt für Mineralogie, Geologie und Paläontologie. 1914, 154-158.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien. Zeitschrift für Induktive Abstammungsund Vererbungslehre. 24, 159-163.
Huene, 1921. Coelurosaurier-Reste aus dem obersten Keuper von Halberstadt. Centralblatt für Mineralogie, Geologie und Paläontologie. 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. VEB Gustav Fischer Verlang, Jena. 1-716.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press, Chicago. 468 pp.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 280-305.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.

Lepidus Nesbitt and Ezcurra, 2015
L. praecisio Nesbitt and Ezcurra, 2015
Early Norian, Late Triassic
Dockum Site 7 General TMM 41936, Colorado City Formation of the Dockum Group, Texas, US
Holotype- (TMM 41936-1.3) distal tibia, distal fibula, astragalocalcaneum
Referred- ?(TMM 41936-1) femoral fragment (~178 mm) (Nesbitt and Ezcurra, 2015)
?(TMM 41936-1.1) partial maxilla (Nesbitt and Ezcurra, 2015)
Diagnosis- (after Nesbitt and Ezcurra, 2015) well-developed posterior pyramidal process on astragalus that delimits posterolateral margin of tibial facet and posteromedial portion of fibular facet, and is separated from proximal surface of calcaneum by shallow notch that opens dorsolaterally.
Comments- The material was discovered in 1941, but not described and named until 2015. The referred material was found in the same area, is congruent in size with the holotype, and matches expected coelophysoid morphology. Using a version of Nesbitt's basal dinosauromorph matrix, Lepidus emerged as a coelophysid closer to Coelophysis than kayentakatae when only the holotype was coded (or other places in Neotheropoda with one extra step). When the referred material was included, Lepidus was a coelophysoid outside kayentakatae+Coelophysis. Wang et al. (2017) used another analysis to recover Lepidus sister to Coelophysis bauri within Coelophysidae.
References- Nesbitt and Ezcurra, 2015. The early fossil record of dinosaurs in North America: A new neotheropod from the base of the Upper Triassic Dockum Group of Texas. Acta Palaeontologica Polonica. 60(3), 513-526.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.

Longosaurus Welles, 1984
L. longicollis Welles 1984
Late Norian, Late Triassic
Arroyo Seco, Petrified Forest Member of the Chinle Formation, New Mexico, US
Holotype- (AMNH 2705; = AMNH 2701 before 1973; paralectotype of Coelurus longicollis) partial ilium
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. No features are obviously more similar to coelophysoids than to Liliensternus, making this Neotheropoda indet..
Reference- Cope, 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proceedings of the American Philosophical Society. 24(126), 209-228.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.

"Megalosaurus" woodwardi Lydekker, 1909
= Megalosaurus lydekkeri Huene, 1926
= Magnosaurus lydekkeri (Huene, 1926) Huene, 1932
Sinemurian, Early Jurassic
Lower Lias, England
Holotype- (NHMUK 41352) maxillary or dentary fragment, tooth (25 x 10 x ? mm)
Comments- Note the name Megalosaurus woodwardi has been independently created two other times. Once by Huene (1932) as a (probably accidentally retained) assignment of his new species Magnosaurus woodwardi (itself an objective junior synonym of Sarcosaurus adrewsi), and again by Olshevsky (1991) as an unattributed supposed junior synonym of Walgettosuchus woodwardi. These are all based on different holotypes, though ironically Lydekker (1909) did consider the Sarcosaurus andrewsi material to belong to Megalosaurus woodwardi.
This tooth was first mentioned by Dawkins (in Huxley, 1869) as Megalosaurus. Purchased by the NHMUK in 1869, Lydekker (1888) described it as Zanclodon(?) sp. b. as he felt the large degree of labiolingual compression and longitudinal striations were more similar to that genus than to Megalosaurus. Lydekker (1909) reviewed Woodward's (1908) paper describing the tibia that would later be named Sarcosaurus andrewsi, and felt the tibia and tooth belonged to the same species, which he named Megalosaurus woodwardi, designating the tooth as the type. Huene (1926) was apparently unaware of the 1909 paper and named NHMUK 41352 Megalosaurus (gen. ?) lydekkeri, stating simple curvature distinguished it from Megalosaurus terquemi. In 1932, Huene reassigned the species to Magnosaurus, again with reservations, though without stating his reason. He believed unspecified remains from the Lower Lias of Watchet referred to Megalosaurus by Phillips might belong to the same species, but as these have never been described this is uncertain. NHMUK 41352 has since been called Megalosaurus or Magnosaurus lydekkeri and is usually placed as Theropoda indet., as in the most recent reviews by Benson and Barrett (2009) and Carrano et al. (2012). Carrano et al. considered it likely to be non-tetanurine as it's striated and "the tooth lacks the specialized features of tetanurans in which such striations also occur (e.g. spinosaurids)." But the striations in M. woodwardi are much narrower and less prominent than the fluting in spinosaurids, and there's no reason they couldn't evolve in another tetanurine lineage.
Because Megalosaurus woodwardi has priority over Megalosaurus lydekkeri, but has not been used since it was named, ICZN Article 23.9.1 should be consulted to check if the former is a nomen oblitum. Usage of Megalosaurus lydekkeri must be maintained if (23.9.1.1) M. woodwardi has not been used as a valid name after 1899 (false, as Lydekker used it in 1909) and (23.9.1.2) M. lydekkeri has been used as a valid name "in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years" (false as far as I can tell, as an extensive search located only 13-15 works since 1962). Thus Megalosaurus woodwardi is the valid name, unless a worker were to petition the ICZN to suppress it.
References- Huxley, 1869. On the upper jaw of Megalosaurus. Quarterly Journal of the Geological Society of London. 25, 311-314.
Phillips, 1871. Geology of Oxford and the Valley of the Thames: Oxford at the Clarendon Press. 523 pp.
Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamta, Rhynchocephalia, and Proterosauria. British Museum of Natural History, London. 309 pp.
Lydekker, 1909. Vertebrate paleontology in 1908. Science Progress in the Twentieth Century: A Quarterly Journal of Scientific Work & Thought. 3(11), 450-471.
Huene, 1926 The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista Museo de La Plata, 29, 35-167.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte [The fossil reptile order Saurischia, their development and history]. Monographien zur Geologie und Palaeontologie, serie 1. 4(1-2), 1-361.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Benson and Barrett, 2009. Dinosaurs of Dorset: Part I, the carnivorous dinosaurs (Saurischia, Theropoda). Proceedings of the Dorset Natural History and Archaeological Society. 130, 133-147.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

"Newtonsaurus" Welles vide Welles and Pickering, 1999
"N." cambrensis (Newton, 1899) Welles vide Welles and Pickering, 1999
= Zanclodon cambrensis Newton, 1899
= Megalosaurus cambrensis (Newton, 1899) Molnar, Kurzanov and Dong, 1990
= Gresslyosaurus cambrensis (Newton, 1899) Olshevsky, 1991
Rhaetian, Late Triassic
Rhaetic Beds, Wales
Holotype- dentary (275 mm), teeth (to 29 mm)
Comments- Rauhut and Hungerbuhler (2000) note that the three supposed derived characters shared with Megalosaurus are not valid. The angular rostral margin is found in Liliensternus, Syntarsus and Sellosaurus for instance. The separate interdental plates are found in Plateosaurus, Dilophosaurus and several other theropods. The third character, "replacement teeth exposed at base between interdental plates", is correlated with separate interdental plates. The authors find it agrees quite well with Liliensternus and Dilophosaurus, but refer it to Theropoda indet.. Welles (1984) found several differences from Dilophosaurus, so I think we should wait for an in depth analysis to proclaim this specimen indeterminate. If added to the matrix of Carrano et al. (2012), it emerges as a noasaurid, suggesting it may be a basal ceratosaur, though the authors considered it a non-averostran.
References- Newton, 1899. On a megalosauroid jaw from Rhaetic Beds near Bridgend (Glamorganshire). Quarterly Journal of the Geological Society of London. 55, 89-96.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. Berkeley: University of California Press. 169-209.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Welles and Pickering, 1999. An Extract From: Archosauromorpha: Cladistics and Osteologies. 70 pp.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

Protoaviformes Chatterjee, 1991
= Praeornithurae Kurochkin, 1995
= Protoaviornithes Kurochkin, 1995
Comments- Chatterjee (1991) named Protoaviformes as a monotypic taxon within his concept of Aves. Kurochkin (1995) named Praeornithurae as a subclass of his Aves, only containing the infraclass Protoaviornithes.
Protoavidae Chatterjee, 1991
Protoavis Chatterjee, 1991
= "Protoavis" Chatterjee vide Wilford, 1986
P. texensis Chatterjee, 1991
Middle Norian, Late Triassic
Post Quarry MOTT 3624, Lower Cooper Canyon Formation of the Dockum Group, Texas, US
Holotype- (TTU-P9200)
unassigned- premaxilla, maxilla, lacrimal(?), posterior mandible(?), atlantal intercentrum, posterior cervical vertebra (18 mm), ?second caudal vertebra (10 mm), caudal vertebra, incomplete clavicle(?), proximal scapula(?), ilia(?) (one incomplete, one partial; ~43 mm), pubis(?) (37 mm), partial ischia(?) (22 mm), distal femur, phalanx I-1(?) (17 mm), phalanx II-1(?) (12 mm), phalanx II-2(?) (11 mm), pedal ungual II(?) (11 mm), phalanx III-1(?) (14 mm), pedal ungual III(?) (5 mm)
Simiosauria- squamosal, quadrate, frontal, parietal, axis (10 mm)
Pterosauromorpha- ischium
Neotheropoda- (juvenile?) basioccipital, prootic, exoccipital, epiotic, supraoccipital, proximal femur
Paratype- (TTU-P9201)
unassigned- maxilla, nasal(?), lacrimal, jugal, quadratojugal, quadrate(?), parietal(?), basioccipital(?), vomer, palatine, pterygoid, partial dentary, posterior dentary(?), posterior mandible, cervical rib, seven dorsal vertebrae (10, 8, 8, 8, 8, 7, 7 mm), two dorsal ribs, two sacral vertebrae (7 mm), eighteen caudal vertebrae (8, 7, 9, 9, 8, 7, 8, 9, 8, 8, 9, 8, 7, 7, 8, 8, 8, 8 mm), two chevrons, incomplete scapula(?), coracoid(?) (18 mm), incomplete humerus, incomplete ilium(?) (~22 mm), proximal tibia, fibulae (one proximal; 50 mm), metatarsal I(?) (9 mm), phalanx I-1(?) (10 mm)
Simiosauria- axis (8 mm), four cervical vertebrae (10, 12, 11, 12 mm)
Archosauromorpha- posterior cervical vertebra (9 mm), two incomplete posterior cervical vertebrae
Pterosauromorpha- incomplete tibia, incomplete fibula, four tarsals, metatarsal I (12 mm), incomplete phalanx I-1, incomplete pedal ungual I, metatarsal II (20 mm), phalanx II-1, phalanx II-2, incomplete pedal ungual II, metatarsal III (18 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (14 mm),
Neotheropoda- (juvenile?) astragalus, calcaneum, distal tarsal IV, metatarsal II (25 mm), metatarsal III (27 mm), metatarsal IV (25 mm)
Early Norian, Late Triassic
Kirkpatrick Quarry MOTT 3628, Lower Cooper Canyon Formation of the Dockum Group, Texas, US
Referred- (TTU-P9350) dorsal vertebra (10 mm) (Chatterjee, 1999)
(TTU-P9351) dorsal centrum (8 mm) (Chatterjee, 1999)
(TTU-P9352) dorsal vertebra (6 mm) (Chatterjee, 1999)
(TTU-P9353) dorsal centrum (6 mm) (Chatterjee, 1999)
(TTU-P9354) dorsal vertebra (6 mm) (Chatterjee, 1999)
(TTU-P9355) dorsal vertebra (10 mm) (Chatterjee, 1999)
(TTU-P9356) caudal vertebra (9 mm) (Chatterjee, 1999)
(TTU-P9357) caudal vertebra (9 mm) (Chatterjee, 1999)
(TTU-P9358) caudal vertebra (9 mm) (Chatterjee, 1999)
(TTU-P9359) caudal vertebra (8 mm) (Chatterjee, 1999)
(TTU-P9360) proximal coracoid(?) (Chatterjee, 1999)
(TTU-P9361) incomplete sternum(?) (Chatterjee, 1998)
(TTU-P9362) partial humerus (Chatterjee, 1991)
(TTU-P9263) proximal humerus (Chatterjee, 1998)
(TTU-P9364) partial mandible (Chatterjee, 1999)
(TTU-P9365) proximal humerus (Chatterjee, 1999)
(TTU-P9367) proximal radius(?) (Chatterjee, 1999)
(TTU-P9368) incomplete radius(?) (Chatterjee, 1999)
(TTU-P9369) (Clevosaurus?) proximal ulna (Chatterjee, 1999)
(TTU-P9370) incomplete femur (Chatterjee, 1999)
(TTU-P9371) distal femur (Chatterjee, 1999)
(TTU-P9372) distal femur (Chatterjee, 1999)
(TTU-P9373) distal femur (Chatterjee, 1999)
(TTU-P9374) distal tibia (Chatterjee, 1999)
(TTU-P9375) pedal phalanx IV-1(?) (10 mm) (Chatterjee, 1999)
(TTU-P9376) pedal ungual IV(?) (12 mm) (Chatterjee, 1999)
(TTU-P9377) pedal ungual I(?) (9 mm) (Chatterjee, 1999)
(TTU-P9378) pedal ungual I(?) (8 mm) (Chatterjee, 1999)
(TTU-P9379) pedal ungual III(?) (~15 mm) (Chatterjee, 1999)
(TTU-P9380) pedal ungual (9 mm) (Chatterjee, 1999)
Diagnosis- A chimaerical taxon whose holotype includes portions of multiple organisms.
Comments- Discovered in Summer 1983 and initially identified as "a juvenile specimen referrable to Coelophysis" (Chatterjee, 1986), the type material was then reported as two individuals of the oldest bird in the popular press (Wilford, 1986). Wilford stated Chatterjee "said he was naming the creature Protoavis, or first bird", with the avian identification based on both characters now recognized in even coelophysid theropods (e.g. large orbits, furcula, sternum, hollow bones) and some generally limited to coelurosaurian clades (large braincase, teeth limited to anterior jaws, keeled sternum, ulnar and metacarpal quill knobs). Beardsley (1986) reported additional bird-like characters claimed by Chatterjee- incomplete postorbital bar, "typically avian" quadrate, elongated forelimbs, "flight muscle attachment sites on the humerus" and (lateral) cnemial crest. Chatterjee (1987) reported thirty cranial characterts shared by Protoavis and Aves but not Archaeopteryx, and the following year suggested a semilunate carpal articulating with a single metacarpal also supported this (Chatterjee, 1988). This relationship 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. This latter idea was based on the combination of primitive characters (low astragalar ascending process; supposed tetradactyl pes which is contradicted by the short metatarsal I identified in the holotype) and bird-like characters of Herrerasaurus that were not explicitly said to be shared with Protoavis (narrow scapula that is more developed in than Protoavis- Elongation Index ~9 vs. 5.8; unspecified pectoral joint similarity; proximally extensive humeral ectocondyle; acetabular antitrochanteric surface that was never proposed in Protoavis; opisthopubic pelvis). Paul also claimed supposed herrerasaur Alwalkeria "shares some distinctive features with "Protoavis."", but the only potential feature mentioned is serrationless teeth which are part of the crocodylomorph skull in the Alwalkeria type, not the dinosaurian portion. Given that celophysoids have a narrower scapula than Protoavis (EI 7.7) and proximally extensive ectocondyle (e.g. rhodesiensis QG/1; contra Paul), only the opisthopubic pelvis remains to connect Protoavis to herrerasaurines. Given the otherwise highly dissimilar pelvic anatomy it is unlikely the supposed pelvis belongs to a herrerasaurian.
Chatterjee (1991) officially named the taxon and described the skull in depth while figuring some of the type postcrania, then described the type and Kirkpatrick postcrania (Chatterjee, 1999). Unfortunately, the material has a highly fractured surface and was repaired and reconstructed prior to being figured, with photographs not indicating clearly which features are real and illustrations being either simplistic (1991) or idealized schematics (1995-1999). Notably the appendicular measurements given in 1999 are based on Chatterjee's reconstructions that usually involve unknown lengths of missing shaft, so are not repeated above. Some authors agreed it was avialan (Kurochkin, 1992; Peters, 1994; Kurochkin, 1995; Martin, 2004). When analyzed as a bird naively accepting Chatterjee's interpretations, Protoavis is resolved as- an avialan more derived than Archaeopteryx 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 euornithine. As detailed on this site, almost every supposed bird-like character proposed in Chatterjee's 1991 cranial analysis is also present even in basal theropods (e.g. crista prootica covers anterior temporal recess; middle cerebral vein; cerebellar fossa extends to supraoccipital), seemingly absent in Protoavis (naris posteriorly placed; reduced dorsal maxilla process; dorsal quadratojugal process absent; quadratojugal cotyla on quadrate; pterygoid condyle on qudrate; small olfactory lobes; etc.) or poorly constructed. The same cranial characters are largely repeated in his 1999 analysis, while any discussion of his postcranial characters is marred by the questionable identification of elements' identities or even referral to the same taxon. Note the Kirkpatrick quarry and unassigned type element identifications above listed with question marks are based on Chatterjee's identifications, so could easily be different elements.
Thus most authors have dismissed an avialan identity (Kurochkin, 1991; Ostrom, 1991; Wellnhofer, 1992; Chiappe, 1995; Feduccia, 1996; Ostrom, 1996; Sereno, 1997; Hunt et al., 1998; Renesto, 2000; Nesbitt et al., 2005; Martz et al., 2013). These authors have usually suggested the remains are chimaerical, including a non-avian coelurosaur braincase (Witmer, 2001), simiosaur 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 simiosaurs (Ancistrorhynchus, Avicranium, Dolabrosaurus, Drepanosaurus, Skybalonyx, Harris and Downs, 2002, multiple TTU specimens in Mueller and Chatterjee in prep. in Martz, 2008, Renesto et al., 2009, Martz et al., 2012), lepidosauromorphs (Clevosaurus), coelophysoids (Camposaurus, Coelophysis, "Comanchesaurus", Gojirasaurus, Lepidus) and crurotarsans (phytosaurs, revueltosaurs, aetosaurs, poposauroids, teratosaurids, crocodylomorphs) 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). The inability to trust Chatterjee's identifications has led to the remains being basically ignored over the last decade.
A coelurosaur braincase? Witmer (2001) described the large floccular fossa, cranial pneumatic recesses and metotic strut as coelurosaurian characters in the holotype braincase. However, the metotic strut is present in Dilophosaurus, Notatessaraeraptor and other taxa closer to averostrans than coelophysoids, and comparing Protoavis to "Megapnosaurus" kayentakatae suggests it's possible the supposed anterior opening of the posterior tympanic recess is the actual vagus nerve, while the supposed vagus foramen could be the larger of two hypoglossal (XII) foramina giving it the coelophysoid condition. Anterior temporal recesses go back at least to basal dinosauromorphs, dorsal tympanic recesses are present in coelophysoids (Coelophysis, "Megapnosaurus" kayentakatae) and the latter species even shows a posterior tympanic recess albeit with a smaller and more dorsally located opening than Protoavis. The floccular fossa is no larger than recently figured in kayentakatae or Dilophosaurus. Thus the supposed coelurosaurian characters have been discovered to have a much broader distribution, and even if the cranial foramina of Protoavis were correctly identified they could work for e.g. a juvenile Gojirasaurus if the taxon is close to Notatessaraeraptor.
Simiosaur cervicals? Renesto (2000) noted some Protoavis cervical vertebrae share several characters with Megalancosaurus- "the same general outline", "the prezygapophyses have a convex surface and are vertically oriented", "The presence of "hypapophyses" and of elongate, narrow centra that are concave anteriorly and convex posteriorly, together with low neural spines." Indeed, the hypapophyses of both forms an elongate keel, the posterior central surface is dorsally angled, the neural spine is anteriorly limited, and three dimensionally preserved simiosaur cervical AUP 11362 (Renesto and Fraser, 2003) shows that like Protoavis the central articulations are actually heterocoelous. Note this only applies to what Chatterjee describes as cervicals 2-5, as the cervicals described as 10-12 are more generalized amphicoelous archosauromorph in structure. Renesto also suggested cranial similarities, the skulls of both being "narrow and pointed anteriorly with an inflated ... postorbital region", "with a ventrally bent anterior portion" of the mandible. However, the supposed dentary fragments of Protoavis are ambiguous and too fragmentary to determine curvature, and the general triangular shape is common in amniotes and not even that developed in the rounded premaxilla assigned to the Protoavis holotype. However, the L-shaped frontals with narrow interorbital space and highly offset postorbital processes, broad parietals and anteriorly projecting spurs ("zygomatic process" of Chatterjee) are similar and suggest the holotype skull roof is also simiosaurian. Of Chinle and Dockum simiosaurs, only Avicranium and isolated elements MNA.V.3652 preserve cranial or cervical material. Despite some general similarity to Protoavis material in having a triangular skull with large orbits, expanded endocranium, toothless maxilla and elongate retroarticular process, Avicranium strongly differs from the Protoavis skull roof in having anteriorly broad frontals, ventrally extensive squamosal and dorsally broad quadrate. Although the cervicals of Avicranium are not described or figured well, the neural spines are tall unlike Protoavis, while isolated Chinle cervical centrum MNA.V.3652 is truly procoelous with a tall anterior surface and ventrally angled posterior surface. As more basal Vallesaurus also has anteriorly broad frontals, this may indicate the Protoavis skull roof is most closely related to Megalancosaurus among simiosaurs.
Lepidosauromorph humerus? Witmer (2001) stated "Although the distal humeral condyles indeed are well developed, the humerus compares tolerably well with that of extant squamates, and a lepidosauromorph identification is worthy of consideration." Compared to the contemporaneous Clevosaurus, the deltopectoral crest is developed more distally and laterally, the ectocondyle exists and is especially exposed posteriorly, and entepicondylar and ectepicondylar foramina are absent. These same differences exist with the Middle Triassic Fraxinasaura and Megachirella and the Early Jurassic Gephyrosaurus, suggesting the paratype humerus is not lepidosauromorph.
Coelophysoid hindlimbs? Hutchinson (2001) stated "Chatterjee (1997, 1998) reconstructed the femur of 'Protoavis' with a medially offset femoral head. I have examined TTUP 9200; it appears to be the femur of a small (possibly juvenile) basal theropod. The middle of the femoral shaft is missing and thus the femoral head orientation is not certain. However, the preserved femur is identical to coelophysoid femora ..., and there is no independent evidence that the femoral head was oriented other than craniomedially as in other basal theropods." Nesbitt et al. (2007) in turn wrote it "exhibits the following dinosaur and theropod characters: offset femoral head, ligament sulcus, strongly developed facies articularis antitrochanterica of the femur (from Langer 2004), anterior trochanter with strong trochanteric shelf and a small posterior trochanter. Notably the posterolateral lip on the greater trochanter ("posterior trochanter") and obturator ridge are similar to coelophysoids like "Megapnosaurus" kayentakatae and Coelophysis rhodesiensis, but not herrerasaurids or Dilophosaurus, and the Chindesaurus+Tawa clade lacks a prominent ligament sulcus. Thus the proximal femur is likely to be coelophysoid, and as the proximolateral lip, obturator ridge and trochanteric shelf are only present in robust individuals, this may represent a new tiny taxon instead of a juvenile.
Witmer (2001) wrote the paratype proximal tarsals "clearly derive from a small dinosaur and probably can be assigned to Theropoda, but they show no real avian or even coelurosaur apomorphies. The ascending process of the astragalus is very low. The calcaneus is quite large and shares with the astragalus a very large fibular articular surface" and that he "can see no evidence of the fusion noted by Chatterjee." Nesbitt et al. (2007) further noted the tibial facet is limited to the astragalus (incorrectly described, but correctly referenced to Rauhut's 2003 character 219) and "the astragalus and calcaneum articulated directly distal to the tibia and fibula", both of which are unlike averostrans. The lack of even a rudimentary calcanear tuber is unlike Eoraptor, Herrerasaurus and Eodromaeus, while the non-gluteaform shape is unlike the Chindesaurus+Tawa clade.
Additional Coelophysis-like material is known from the Cooper Canyon Formation (ilium TTU-P10071, tibiae TTU-P11044, TTU-P14786 and TTU-P10534), which may belong to the same taxon
Other material- Many elements of Protoavis appear to be from a new supposed pterosauromorph named and described in a thesis (Atanassov, 2001, 2002), "Procoelous vertebrate taxon A" of Martz et al. (2012). The holotype (Chatterjee, 1991; reidentified as a vomer and pterygoid in 1999) sternum is an ischium, the paratype radius and ulna are the tibia and fibula respectively, and the paratype metacarpus is the metatarsus. The latter was foreseen by Sereno (1997), who wrote "The four-digit manus (Chatterjee 1995), for example, is more appropriately identified as an archosaurian pes." 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 referred vertebrae are from a tanystropheid, simiosaur or other taxon.
The articulated series of paratype posterior cervicals is not similar to simiosaurs and instead resembles basal archosauromorphs in being amphicoelous with offset centrum faces while lacking laminae often present in e.g. dinosaurs and other archosaurian subgroups.
Kirkpatrick material- Chatterjee (1991) notes "A few disarticulated bird bones were found on Collier's ranch", stating that "A beautiful humerus from this quarry shows all the avian hallmarks. So far, only vertebrae, coracoid and limb elements of birds have been recovered from this site, and the affinity of this material will be discussed in a separate paper." He later (1995) states "thirty disarticulated postcranial elements of Protoavis were recovered from a small mound in the Kirkpatrick quarry", and similarly in 1997 "I collected a total of thirty-one isolated postcranial elements of Protoavis from the Kirkpatrick quarry." Both works include composites using elements from both quarries, notably the large-keeled supposed sternum, although no specimen numbers are mentioned or other elements purely based on Kirkpatrick material. His 1998 paper figures the supposed sternum TTU-P9361, humeri TTU-P9362 (the one mentioned in 1991) and TTU-P9363, although it does not mention the Kirkpatrick quarry or specimens in the text. Chatterjee (1999) figures much of the material, but mostly as idealized reconstructions, with only the sternum and humeri from the 1998 paper, supposed radius TTU-P9368 and distal femur TTU-P9370 photographed. The proximal humerus TTU-P9365 is mislabeled TTU-P9364 in Chatterjee's figure 14e-f. Incomplete humerus TTU-P9362 shows the same distal morphology as the paratype with some proportional differences (more flared, entocondyle larger anteriorly and more transversely expanded posteriorly), suggesting it is a related taxon to whatever the poaratype humerus belongs to. Notably, the proximal ulna TTU-P9369 does not resemble the paratype's proximal 'ulna' in having a projected olecranon and expanded shaft, and the possible radius TTU-P9368 is much more slender than the paratype's proximal 'radius', so these elements don't seem to belong to the pterosauromorph tibia and fibula and may be an actual ulna and radius. The ulna strongly resembles Clevosaurus AUP 11178 in general shape in side view, so may be referrable to that taxon. Elzanowski (2008) stated "the Triassic Tecovas formation femoral fragments which were assigned by Chatterjee (1991) to Protoavis, but are better comparable to lizards than to either Protoavis holotype or any birds" are unlike non-bird theropods in lacking an ectocondylar tuber. This refers to the Kirkpatrick femora, and indeed TTU-P9370 differs from the holotype in lacking an ectepicondylar tuber so is from a different taxon. Note the tuber in the holotype does not necessarily indicate it is from a theropod, as they are found in other taxa such as Fruitadens and Dromomeron as well.
References- Beardsley, 1986. Fossil bird shakes evolutionary hypotheses. Nature. 322, 677.
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. 139-150.
Wilford, 1986. Texas fossil may be birds' oldest ancestor. The New York Times. 8-14-1986, section A:1.
Chatterjee, 1987. Skull of Protoavis and early evolution of birds. Journal of Vertebrate Paleontology. 7(3), 14A.
Ostrom, 1987. Protoavis, a Triassic bird? Archaeopteryx. 5, 113-114.
Wellnhofer, 1987. Der älteste fossile Vogel gefunden? Naturwissenschaftliche Rundschau. 40(4), 145.
Chatterjee, 1988. Functional significance of the semilunate carpal in archosaurs and birds. Journal of Vertebrate Paleontology. 8(3) 11A.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster. 464 pp.
Anonymous, 1991. Early bird born late. Nature. 51(6329), 677-678.
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. Priroda. 1991, 43-53.
Chatterjee, 1992. Texas fossil identified as the world's oldest known bird. Journal of the Geological Society of India. 39, 89-90.
Kurochkin, 1992. The oldest bird. Priroda. 1992, 100-101.
Viohl, 1992. No evidence of Triassic birds. Archaeopteryx. 10, 77-79.
Wellnhofer, 1992. Protoavis: The oldest bird? Naturwissenschaftliche Rundschau. 45(3), 107-108.
Chatterjee, 1994. Protoavis from the Triassic of Texas: The oldest bird. Journal für Ornithologie. 135, 330.
Peters, 1994. Die Entstehung der Vogel-Verandern die jungsten Fossilfunde das Modell? In Gutmann, Mollenhauer and Peters (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, Lucas, Heckert, Sullivan and 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. Two new archosaur reptiles from the Late Triassic of Texas. PhD Thesis, Texas Tech University. 352 pp.
Harris and Downs, 2002. A drepanosaurid pectoral girdle from the Ghost Ranch (Whitaker) Coelophysis Quarry (Chinle Group, Rock Point Formation, Rhaetian), New Mexico. Journal of Vertebrate Paleontology. 22(1), 70-75.
Renesto and Fraser, 2003. Drepanosaurid (Reptilia: Diapsida) remains from a Late Triassic fissure infilling at Cromwell Quarry (Avon, Great Britain). Journal of Vertebrate Paleontology. 23(3), 703-705.
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. Journal of Vertebrate Paleontology. 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.
Elzanowski, 2008. The avian femur: Morphology and terminology of the lateral condyle. Oryctos. 7, 1-5.
Martz, 2008. Lithostratigraphy, chemostratigraphy, and vertebrate biostratigraphy of the Dockum Group (Upper Triassic), of southern Garza County, west Texas. PhD thesis, Texas Tech University. 504 pp.
Renesto, Spielmann and Lucas, 2009. The oldest record of drepanosaurids (Reptilia, Diapsida) from the Late Triassic (Adamanian Placerias Quarry, Arizona, USA) and the stratigraphic range of the Drepanosauridae. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 252, 315-325.
Martz, Mueller, Nesbitt, Stocker, Parker, Atanassov, Fraser, Weinbaum and Lehane, 2012. A taxonomic and biostratigraphic re-evaluation of the Post Quarry vertebrate assemblage from the Cooper Canyon Formation (Dockum Group, Upper Triassic) of southern Garza County, western Texas. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103, 1-26.

Velocipes Huene, 1932
V. guerichi Huene, 1932
Middle-Late Triassic
Kocury, Lissauer Breccia, Poland
Holotype- (GPIM UH No. 252) proximal fibula (~300-350 mm)
Diagnosis- (after Skawiński et al., 2017) indeterminate relative to individual/ontogenetic variation expected in Dracoraptor, Liliensternus and Dilophosaurus.
Comments- Note the species was originally spelled Velocipes gürichi by Huene, but this is considered an incorrect original spelling that must be corrected by ICZN Article 32.5.2.1 ("In the case of a diacritic or other mark, the mark concerned is deleted, except that in a name published before 1985 and based upon a German word, the umlaut sign is deleted from a vowel and the letter "e" is to be inserted after that vowel") to result in V. guerichi instead.
Huene (1932) referred this to Podokesauridae as the "size and structure" fit that family best, and when describing Liliensternus stated (translated) "Due to the age difference and the different curvature, the species is certainly different, but it is possible that the genus is the same" (Huene, 1934). Welles (1984) agreed regarding the potential synonymy but believed "Our knowledge of theropod fibulae is so limited that we cannot identify this fibula, and must consider Velocipes gurichi a nomen vanum." Norman (1990) similarly stated "The material is indeterminate" at the level of Theropoda and even identified the element as the "Broken proximal end of a tibia (?fibula)", and Rauhut and Hungerbuhler (2000) concurred "even its identification as a fibula may be doubted" and gave it the ridiculously broad status of "Vertebrata, nomen dubium." Czepiński et al. (2014) reported in an abstract that Huene was basically right all along, identifying it as the proximal fibula of a "probable non-tetanuran neotheropod." This was elaborated on in the publication of Skawiński et al. (2017) that redescribed the taxon in depth and provided detailed comparisons which distinguished it from most saurischians except non-averostran neotheropods. Of the latter, they reported "Dilophosaurus is very similar to Velocipes in all aspects (Welles 1984). There are subtle differences in size and position of M. iliofibularis tubercle, which is positioned slightly more proximally and smaller (45 mm, i.e. 9% of total length of fibula; compared with 52 mm in Velocipes, i.e. 15-17% of total length of fibula) in Dilophosaurus. Fibulae of Liliensternus, Dracoraptor and Velocipes are virtually indistinguishable in shape at first glance... However, Liliensternus lacks well visible M. iliofibularis attachment." However, the authors noted that "is possible that one of the most striking features of Velocipes, the prominence of crest for the insertion of M. iliofibularis, is a result of intraspecific variation, as it occurs in many archosaurs" and that the lack of a proximomedial fibular ridge (unlike older coelophysoids) may be due to being ontogenetically young. Given this potential intraspecific variation, Skawiński et al. conclude "probably it is not possible to identify Velocipes more precisely than Theropoda indet.", but this should be revised to Neotheropoda indet. as they do distinguish it from herrerasaurids, Eoraptor, Tawa and Eodromaeus.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Huene, 1934. Ein neuer Coelurosaurier in der thüringischen Trias. Paläontologische Zeitschrift. 16(3/4), 145-170.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 280-305.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia 15, 75-88.
Czepiński, Niedźwiedzki, Tałanda, Skawiński, Ziegler and Szermański, 2014. A re-evaluation of the purported dinosaur finds from the Middle-Late Triassic of Poland. Journal of Vertebrate Paleontology, Program and Abstracts, 2014. 115.
Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017 (online 2016). A re-evaluation of the historical 'dinosaur' remains from the Middle-Upper Triassic of Poland. Historical Biology. 29(4), 442-472.

Coelophysoidea Nopcsa, 1928 sensu Holtz, 1994
Definition- (Coelophysis bauri <- Carnotaurus sastrei) (modified from Sereno, 1998)
Other definitions- (Coelophysis bauri <- Ceratosaurus nasicornis) (Ezcurra, 2017; modified from Padian et al., 1999)
(Coelophysis bauri <- Ceratosaurus nasicornis, Carnotaurus sastrei, Passer domesticus) (Ezcurra and Brusatte, 2011)
(Coelophysis bauri <- Passer domesticus) (Allain et al., 2012)
(Coelophysis bauri <- Allosaurus fragilis, Ceratosaurus nasicornis) (Dal Sasso, Maganuco and Cau, 2018)
= Podokesauroidea Huene, 1914 vide 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 Ezcurra and Brusatte, 2011
(Coelophysis bauri <- Ceratosaurus nasicornis, Carnotaurus sastrei, Passer domesticus)
= Coelophysoidea sensu Dal Sasso, Maganuco and Cau, 2018
Definition- (Coelophysis bauri <- Allosaurus fragilis, Ceratosaurus nasicornis)
Diagnosis- (suggested) infrapopliteal ridge on femur; medial edge of distal tarsal IV flat (absent in Panguraptor); metatarsals II and III fused proximally (ontogenetic); in proximal view, metatarsal IV does not extend posterior to III (absent in Panguraptor).
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 nomenclaturally valid). Coelophysoidea is 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 (e.g. Cryolophosaurus, Dracovenator and Sinosaurus; called dilophosaurs here as they may not be monophyletic) are coelophysoids, or more closely related to averostrans. Traditional phylogenies placed them in Coelophysoidea (see above), and this was found in many 2000s analyses as well (Wilson et al., 2003; Holtz et al., 2004; Tykoski and Rowe, 2004; Tykoski, 2005; Ezcurra and Novas, 2007). Yet a few 2000s analyses instead recovered Dilophosaurus as more closely related to averostrans (Rauhut, 2003; Yates, 2005; Smith et al., 2007). Rauhut found Dilophosaurus to be equally well supported as a coelophysoid when Shuvosaurus was (correctly) excluded, Yates found dilophosaurs were coelophysoids with only one added step, and Smith et al. found dilophosaurs 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 Sinosaurus, whereas Smith et al. included all three and Yates includes Dracovenator. Also relevant is that Tykoski found Dilophosaurus to have been misscored in prior analyses due to a reliance on subadult specimens. All more recent analyses recover Dilophosaurus closer to averostrans than Coelophysis, such as the taxonomically extensive Wang et al. (2016) which takes eighteen more steps to place Dilophosaurus in Coelophysoidea. Cau's megamatrix has been published in several analyses, with Dal Sasso et al. (2018) focusing on basal neotheropods where it takes only two extra steps to place Dilophosaurus in Coelophysoidea (as happened using the different taxon set in Cau, 2018). Most recent coelophysoid-grade taxa have been described using versions of Nesbitt's dinosauromorph analysis which recover Dilophosaurus closer to averostrans and have Ezcurra et al. (2021) as the most recent incarnation incorporating the most taxa. This topology agrees with Ezcurra's (2012) analysis "composed of 39 terminals and 633 informative characters" which remains unpublished, and the results of correcting some scores in this analysis form the basis of the topology used on this site. Enforcing coelophysoid Dilophosaurus requires only three additional steps. Thus the current popularity of non-coelophysoid Dilophosaurus is not due to strong character support, and much could probably be revealed by comparing the strongly supported coelophysoid Dilophosaurus of Tykoski (2005) with the opposite in Wang et al. (2016).
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 (Benton and Walker, 2010), 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). Maisch and Matzke (2003) tentatively assigned tooth GPIT SGP 2001/5 and supposed distal fibula (actually an ischium?) GPIT SGP 2000/2 from the Qigu Formation of China to Coelophysoidea, but given their Late Jurassic age and lack of convincing synapomorphies they are assigned to Averostra here. 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. Novas et al. (2009) called femur ISI R283 from the Lower Dharmaram Formation of India a coelophysoid, but it was later described less specifically as a non-averostran theropod (Novas et al., 2010). Dracoraptor was found to be a coelophysoid in its description (Martill et al., 2016), but this is based on miscodings, and the taxon actually falls out more basally by Daemonosaurus and Chilesaurus.
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. 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, Abteilung A. 185, 85-180.
Carroll, 1988. Vertebrate Paleontology and Evolution. W.H. Freeman and Company. 698 pp.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 464 pp.
Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9(2), 125-136.
Novas, 1991. Relaciones filogeneticas de los dinosaurios teropodos ceratosaurios. 28(3-4), 401.
Novas, 1992. La evolucion de los dinosaurios carnivoros. 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. Neues Jahrbuch fur Geologie und Palaontologie. 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 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.
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.
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. Contributions from the Museum of Paleontology. The University of Michigan. 31, 1-42.
Heckert, Lucas, Rinehart and Hunt, 2004. Biostratigraphy, biochronology, and evolutionary trends of coelophysoids (Theropoda: Ceratosauria). Journal of Vertebrate Paleontology. 24(3), 206A-207A.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 25-46.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 47-70.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Thesis, 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.
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.
Ezcurra and Novas, 2007 (online 2006). Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
Smith and Makovicky, 2007. Early theropod evolution and paraphyly of the Coelophysoidea. Journal of Vertebrate Paleontology. 27(3), 150A.
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.
Novas, Chatterjee, Ezcurra and Kutty, 2009. New dinosaur remains from the Late Triassic of Central India. Journal of Vertebrate Paleontology. 29(3), 156A.
Benton and Walker, 2010. Saltopus, a dinosauriform from the Upper Triassic of Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 285-299.
Ezcurra and Brusatte, 2011. Taxonomic and phylogenetic reassessment of the early neotheropod dinosaur Camposaurus arizonensis from the Late Triassic of North America. Palaeontology. 54(4), 763-772.
Novas, Ezcurra, Chatterjee and Kutty, 2010. New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of central India. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 333-349.
Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften. 99(5), 369-377.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Martill, Vidovic, Howells and Nudds, 2016. The oldest Jurassic dinosaur: A basal neotheropod from the Hettangian of Great Britain. PLoS ONE. 11(1), e0145713.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
Ezcurra. 2017. A new early coelophysoid neotheropod from the Late Triassic of northwestern Argentina. Ameghiniana. 54, 506-538.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Dal Sasso, Maganuco and Cau, 2018. The oldest ceratosaurian (Dinosauria: Theropoda), from the Lower Jurassic of Italy, sheds light on the evolution of the three-fingered hand of birds. PeerJ. 6:e5976.
Ezcurra, Butler, Maidment, Sansom, Meade and Radley, 2021 (online 2020). A revision of the early neotheropod genus Sarcosaurus from the Early Jurassic (Hettangian-Sinemurian) of central England. Zoological Journal of the Linnean Society. 191(1), 113-149.

unnamed possible ceolophysoid (Jenkins, Foster and Gay, 2017)
Rhaetian, Late Triassic
Corral Canyon, Church Rock Member of the Chinle Formation, Utah, US
Material- (MWC 5627) incomplete synsacrum (s4 26 mm)
Comments- Discovered in 2005, this was described by Jenkins et al. (2017) as a neotheropod based on five sacrals, sacral fusion and subequal centrum lengths, and was said to be "visibly identical to NMMNH 31661", a Snyder Quarry coelophysoid synsacrum. Marsh and Parker (2020) stated it "is likely a coelophysid owing to the co-ossification of the sacral centra."
References- Jenkins, Foster and Gay, 2017. First unambiguous dinosaur specimen from the Upper Triassic Chinle Formation in Utah. Geology of the Intermountain West. 4, 231-242.
Marsh and Parker, 2020. New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils. PaleoBios. 37, 1-56.

Coelophysis? longicollis (Cope, 1887a) Cope, 1889
= Coelurus longicollis Cope, 1887a
= Tanystropheus longicollis (Cope, 1887a) Cope, 1887b
Late Norian, Late Triassic
Arroyo Seco, Petrified Forest Member of the Chinle Formation, New Mexico, US
Lectotype- (AMNH 2701) cervical vertebra (63 mm)
Comments- 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 coelophysoids.
References- Cope, 1887a. The dinosaurian genus Coelurus. American Naturalist. 21, 367-369.
Cope, 1887b. A contribution to the history of the Vertebrata of the Trias of North America. Proceedings of the American Philosophical Society. 24, 209-228.
Cope, 1889. On a new genus of Triassic Dinosauria. American Naturalist. 23, 626.
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.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
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.

Kayentavenator Gay, 2010
K. elysiae Gay, 2010
Sinemurian-Pliensbachian, Early Jurassic
Willow Springs 2 UCMP V82309, Silty Facies Member of the Kayenta Formation, Arizona, US
Holotype- (UCMP V128659; paratype of Syntarsus kayentakatae) (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 bauri and Segisaurus, though seem present in at least some Coelophysis rhodesiensis. 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 Coelophysis rhodesiensis 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 Coelophysis rhodesiensis, 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 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 October 1981 (UCMP online, contra Rowe, 1989) 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 Coelophysis rhodesiensis 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 NHMUK 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 Coelophysis rhodesiensis 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, Coelophysis rhodesiensis) 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 Coelophysis rhodesiensis 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 Coelophysis rhodesiensis, 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. Ezcurra (2012) found Kayentavenator to be sister to kayentakatae and so suggested their synonymy, based on a large unpiblished analysis.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9(2), 125-136.
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications for ceratosaurid phylogeny. Masters Thesis. University of Texas at Austin. 217 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Thesis, University of Texas at Austin. 553 pp.
Gay, 2010. Notes on Early Mesozoic theropods. Lulu Press. 44 pp.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.

Pendraig Spiekman, Ezcurra, Butler, Fraser and Maidment, 2021
P. milnerae Spiekman, Ezcurra, Butler, Fraser and Maidment, 2021
Rhaetian, Late Triassic
Pant-y-ffynnon Quarry, Wales
Holotype- (NHMUK R 37591; field numbers P76/1 and P77/1) (robust subadult) incomplete twelfth dorsal vertebra, incomplete thirteenth dorsal vertebra (14.7 mm), incomplete first sacral vertebra, incomplete fused second-fourth sacral vertebre (13.3,11.1,? mm), ilium (55.8 mm), incomplete pubes, partial ischia, incomplete femur (~102.1 mm)
Paratypes- ?(NHMUK R 37596; field number P83/1) mid-posterior dorsal vertebra (14.6 mm)
(NHMUK R 37597; field number P65/66b) proximal ischium
....(lost; field number P65/66a) ischial fragment
Diagnosis- (after Spiekman et al., 2021) absence of dorsal hyposphene-hypantrum articulations; anteriorly expanded dorsal neural spine; posteriormost dorsal vertebrae with strongly elongated centrum (centrum length ~2.6 times its anterior height) (also in Coelophysis); distinctly anteroventrally slanted dorsal margin of preacetabular process (also in Sarcosaurus); posterodorsal margin of postacetabular process curving abruptly posteroventrally so that posteroventral end is formed by an acute angle of approximately 65 degrees in lateral view; pubis with pubic fenestra (also in coelophysines and Gojirasaurus); ischium with shallow obturator notch; fourth trochanter posteriorly developed to a height similar to the depth of the shaft at that level (also in Procompsognathus, Liliensternus, Dilophosaurus and Cryolophosaurus).
Comments- Initially described as a coelurosaur in Warrener's (1983) thesis, this material was collected between 1951 and 1962. She felt several additional elements were possibly referrable (metapodial field number P65/23, phalanges P65/30 and P65/49 and ungual P65/45), but Spiekman et al. (2021) said "these elements do not exhibit diagnostic theropod features" so they are listed as Archosauriformes here. Fraser and Padian (1995) mention "associated and articulated material of ... Syntarsus." Rauhut and Hungerbuhler (2000) described the holotype specimens as ?Syntarsus sp., and said they "are extremely similar to the coelophysid Syntarsus" and "furthermore very similar to Procompsognathus, and new discoveries might prove that they are referable to this genus." Note these authors used field numbers for the specimens (BMNH PV RU P 77/1 for the pelvis and articulated vertebrae, BMNH PV RU P 76/1 for the femur), whereas Spiekman et al. reported both are catalogued as NHMUK PV R 37591. Galton and Kermack (2010) mention it as "vertebrae and the pelvic girdle and hind limb of the coelurosaurian theropod dinosaur Coelophysis." Although Keeble et al. (2018) stated "We could not locate these fossils in the NHMUK collections", Spiekman et al. say "the articulated partial pelvic girdle and vertebrae, as well as the femur and complete isolated dorsal vertebra referred to the theropod by Warrener have now been relocated" and described the specimens in detail as a new taxon of coelophysoid Pendraig milnerae. Spiekman et al. used Ezcurra's version of Nesbitt's dinosauromorph matrix and recovered it as a non-coelophysid coelophysoid in a polytomy with Lucianovenator, Powellvenator and kayentakatae+Coelophysidae. It is placed in Coelophysoidea here based on the elongate dorsal centra, fused synsacrum and wider angle between the pubic peduncle and preacetabular process.
References- Warrener, 1983. An archosaurian fauna from a Welsh locality. PhD thesis, University College London. 384 pp.
Fraser and Padian, 1995. Possible dinosaur remains from Britain and the diagnosis of the Dinosauria. Journal of Vertebrate Paleontology. 15(3), 30A.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.
Galton and Kermack, 2010. The anatomy of Pantydraco caducus, a very basal sauropodomorph dinosaur from the Rhaetian (Upper Triassic) of South Wales, UK. Revue de Paléobiologie. 29(2), 341-404.
Keeble, Whiteside and Benton, 2018. The terrestrial fauna of the Late Triassic Pant-y-ffynnon Quarry fissures, South Wales, UK and a new species of Clevosaurus (Lepidosauria: Rhynchocephalia). Proceedings of the Geologists' Association. 129(2), 99-119.
Spiekman, Ezcurra, Butler, Fraser and Maidment, 2021. Pendraig milnerae, a new small-sized coelophysoid theropod from the Late Triassic of Wales. Royal Society Open Science. 8: 210915.

Podokesaurus Talbot, 1911
P. holyokensis Talbot, 1911
= Coelophysis holyokensis (Talbot, 1911) Colbert, 1964
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. It is a coelophysoid based on the elongate dorsal centra and may not be a coelophysine based on the anteriorly flexed distal humerus.
References- Talbot, 1911. Podokesaurus* holyokensis, a new dinosaur from the Triassic of the Connecticut Valley. American Journal of Science. 31(186), 469-479.
Colbert and Baird, 1958. Coelurosaur bone casts from the Connecticut Valley Triassic. American Museum Novitates. 1901, 1-11.
Colbert, 1964. The Triassic dinosaur genera Podokesaurus and Coelophysis. American Museum Novitates. 2168, 12 pp.

Pterospondylus Jaekel, 1914
P. trielbae Jaekel, 1914
Early Rhaetian, Late Triassic
Trossingen Formation, Halberstadt, 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 Coelophysis rhodesiensis more than Liliensternus and Dilophosaurus. Thus, Rauhut and Hungerbuhler (2000) suggest the taxon may be a coelophysoid.
References- Jaekel, 1914. Über die Wirbeltierfunde in der oberen Trias von Halberstadt. Paläontologische Zeitschrift. 1, 155-215.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.

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 Averostra); 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 anterior articular surface of proximal caudal vertebrae (also present in Averostra); constant length of caudal vertebrae along the tail (also in Dilophosaurus).
Other diagnoses- 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.
Comments- While often placed close to Liliensternus, the wide notch below the preacetabular process may make this less closely related to averostrans than that 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, 261-288.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis, University of Bristol. 440 pp.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.
Ezcurra and Cuny, 2007. The coelophysoid Lophostropheus airelensis, gen. nov.: A review of the systematics of "Liliensternus" airelensis from the Triassic-Jurassic boundary outcrops of Normandy (France). Journal of Vertebrate Paleontology. 27(1), 73-86.

Powellvenator Ezcurra, 2017
P. podocitus Ezcurra, 2017
Middle Norian, Late Triassic
Los Colorados Formation, La Rioja, Argentina
Holotype- (PVL 4414-1) (subadult) (~700-725 g) distal tibia (10.7 mm wide), distal fibula, astragalus (10.6 mm wide), calcaneum, distal tarsal III fused to distal tarsal IV, proximal metatarsal II fused to proximal metatarsal III, proximal metatarsal IV, incomplete metatarsal V
?...(PVL 4414-3) proximal tibia
?...(PVL 4414-4) proximal tibia, proximal fibula
?...(PVL 4414-5) distal femur (10 mm wide)
?...(PVL 4414-8) distal femur (10.4 mm wide), fragment
Referred- (PVL 3848 in part; lost) femur (~94 mm), incomplete tibia, proximal fibula, astragalus (~15 mm wide) (Bonaparte, 1971)
Diagnosis- (after Ezcurra, 2017) tibia lacking an anterior diagonal tuberosity; tibia lacking a notch for reception of a posteromedial process of the astragalus; astragalus with an ascending process lower than the astragalar body; distinctly sigmoid (medially convex and laterally concave) posterodorsal margin of astragalus; rounded, dorsally projected expansion on the medial third of the dorsal margin of the astragalus in anterior view; calcaneum with laterally projected flange that occupies the entire proximodistal height of its anterolateral corner; distal tarsal III with convex anterolateral margin in proximal view; distal tarsal IV with strongly convex anterior margin in proximal view; shaft of metatarsal II less than half the anteroposterior depth and transverse width of shaft of metatarsal III; metatarsal III without overlap of posteromedial surface of metatarsal IV.
Comments- Note Ezcurra states the non-holotype material is considered referred instead of paratype, "in case that in the future it is shown that they do not belong to the same species as the holotype." The material was discovered in the late 1960s, PVL 4414 "next to the holotype of Riojasuchus" (Ezcurra, 2017). For PVL 4414, Ezcurra writes "the size is congruent for a single neotheropod individual and the character-states are consistent for a single neotheropod taxon." PVL 3848 was originally described by Bonaparte (1971) as Coelurosauria indet., under the Hueneian concept that included coelophysoids. Indeeed, he states Podokesauridae is the "group that best matches the proportions of the bones described" [translated]. Novas (1989) later redescribed and identified it as Theropoda indet. in his thesis. Three partial cervical vertebrae were originally placed in this specimen, but Ezcurra states "these vertebrae are smaller than expected for a single individual in comparison with the hindlimb bones and they do not show neotheropod apomorphies (e.g., pleurocoels). As a result, these three articulated cervical vertebrae were tentatively excluded from PVL 3848." They are placed here tentatively as Saurischia indet.. Ezcurra writes "PVL 3848 could not be located in the collections of the PVL after several visits of the author during the last ten years and should be considered currently lost (J. Powell pers. comm., 2015)."
Ezcurra used a version of Nesbitt's archosaur matrix to recover Powellvenator as a coelophysid closer to coelophysines+segisaurines than kayentakatae and Panguraptor. Yet only two steps place it as a basal coelophysoid or basal neotheropod, and only three steps as sister to averostrans.
References- Bonaparte, 1971. Los tetrapodos del sector superior de la Formacion Los Colorados, La Rioja, Argentina. (Triásico Superior) I parte. Opera Lilloana. 22, 1-183.
Novas, 1989. Los dinosaurios carnívoros de la Argentina. PhD thesis, Museo de La Plata. 510 pp.
Ezcurra, 2017. A new early coelophysoid neotheropod from the Late Triassic of northwestern Argentina. Ameghiniana. 54, 506-538.

Coelophysidae Nopcsa, 1928 vide Welles, 1984
Definition- (Coelophysis bauri + Coelophysis rhodesiensis + "Syntarsus" kayentakatae) (Holtz, 1994)
Other definitions- (Coelophysis bauri + Procompsognathus triassicus) (Sereno, in press; modified from Sereno, 1998)
(Coelophysis bauri + Coelophysis rhodesiensis) (modified from Tykoski and Rowe, 2004)
= Podokesauridae Huene, 1914a
= Podokesaurinae Huene, 1914a vide Nopcsa, 1923
= Procompsognathidae Nopcsa, 1923 vide Huene, 1929
= Segisauridae Camp, 1936
Diagnosis- (suggested) elongate retroarticular process (unknown in Lucianovenator); deep proximomedial fibular fossa; tibia fused to astragalus (ontogenetic); distal tarsal III fused to metatarsal III (ontogenetic).
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 (1914a) 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), Coelophysis rhodesiensis (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 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 (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).
Procompsognathidae- 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.
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, as defined in Tykoski and Rowe (2004), then Segisauridae may be used for its sister clade, including Segisaurus, "Syntarsus" 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, 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 Coelophysis rhodesiensis, 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 bauri and Coelophysis 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 "Syntarsus" kayentakatae. This is far more exclusive than most concepts of Coelophysidae (or Podokesauridae) have been. Also, the synapomorphies for Coelophysis are all cranial, making the assignment of postcrania to Coelophysidae impossible.
References- Huene, 1914a. Beiträge zur geschichte der Archosaurier. Geologie und Paläontologie Abhandlungen. 13(7), 1-56.
Huene, 1914b. Das natürliche System der Saurischia. Centralblatt für Mineralogie, Geologie und Paläontologie. 1914, 154-158.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien. 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. 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. Paläontologische Zeitschrift. 11, 269-273.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of Arizona. University of California Publications in Geological Sciences. 24(2), 39-56.
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. 772 pp.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press, Chicago. 468 pp.
Raath, 1969. A new coelurosaurian dinosaur from the Forest Sandstone of Rhodesia. Arnoldia. 4(28), 1-25..
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp.
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. 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, Abteilung A. 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. 139-150.
Chatterjee, 1987. A new theropod dinosaur from India with remarks on the Gondwana-Laurasia connection in the Late Triassic. Geophysical Monographs. 41, 183-189.
Carroll, 1988. Vertebrate Paleontology and Evolution. W.H. Freeman and Company. 698 pp.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Hunt and Lucas, 1989. Late Triassic vertebrate localities in New Mexico. In Lucas and Hunt (eds.). Dawn of the Age of Dinosaurs in the American Southwest. 72-101.
Novas, 1991. Relaciones filogeneticas de los dinosaurios teropodos ceratosaurios. Ameghiniana. 28(3-4), 401.
Novas, 1992. La evolucion de los dinosaurios carnivoros. 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 pp.
Rauhut and Hungerbuhler, 2000 (as 1998). 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(3), 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. Journal of Vertebrate Palaeontology. 24(3), 117-118.
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 25-46.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Thesis, 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.
Stiegler, Wang, Xu and Clark, 2013. Coding individual specimens as taxa: Test cases aid in resolving the relationships of basal Neotheropoda, gauge topological sensitivity to taxon sampling, and produce novel taxonomic hypotheses. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 219-220.

unnamed coelophysid (Padian, 1986)
Late Norian, Late Triassic
Dinosaur Hill PFV 040 / Inadvertent Hills UCMP V82250, Petrified Forest Member of Chinle Formation, Arizona, US
Material- (PEFO 21373; = UCMP 129618; Lacey Point theropod; Padian theropod; Padian's Coelophysis; Petrified Forest theropod) (subadult) dorsal centrum (45 mm), partial dorsal centrum, two synsacral fragments, proximal caudal centrum (33 mm), ilia (one partial, one incomplete), pubes (one incomplete, one distal), partial ischia, femora (one incomplete; 245 mm), tibiae (270, 255 mm), fibulae (one incomplete; 250 mm), astragalocalcanea, distal tarsal IV, phalanx I-1 (29 mm), distal tarsal III fused to 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)
Late Norian, Late Triassic
Giving Site PFV 231, Petrified Forest Member of Chinle Formation, Arizona, US
?(PEFO 33981) fragmentary skeleton including partial posterior vertebral column, pelvis, incomplete hindlimbs including femora, tibia and astragalocalcaneum (Parker and Irmis, 2005)
(PEFO 33983) incomplete posterior skeleton including proximal femur, tibia, astragalocalcaneum and distal tarsal IV (Parker and Irmis, 2005)
Late Norian, Late Triassic
Jeremiah's Perch PFV 278, Petrified Forest Member of Chinle Formation, Arizona, US
?(PEFO coll.) partial skeleton including teeth, cervical vertebrae, femora and tibiae (Hunt and Wright, 1999)
Comments- Discovered on August 21 1982, UCMP 129618 was described by Padian (1986) as a specimen of Coelophysis bauri. This was because "compared to the type material. they correspond exactly in both size and detail to the largest pieces in Cope's collection", although Padian did state "there are differences between the Ghost Ranch [ = 'Rioarribasaurus', = Coelophysis bauri neotype quarry] and Petrified Forest specimens that may require taxonomic reevaluation when full descriptions of the former are published." Hunt and Lucas (1991) agreed, calling it the Lacey Point theropod "and note several differences including better development of the posteroventral arch of the ilium ... a distinct emargination on the distal tibia for reception of the ascending process of the astragalus ... a much more distinctly offset [femoral] head ... a shorter fourth trochanter ... a less well developed trochanteric shelf [and] a well developed obturator foramen." They concluded "UCMP 129618 represents a ceratosaurian ... distinct from Rioarribasaurus that probably merits a new generic name." Hunt et al. (1998) stated it differed from the Coelophysis neotype in having "(1) femoral head more offset from the femoral shaft, with a deep groove on the proximal face of the head and a concave distal margin to the head so that the medioventral corner of the head forms a ventrally directed point; (2) a tibia with a more robust cnemial crest and very large crest for attachment of the flexor muscle and ligaments on the anterior face; (3) a larger ascending process of astragalus; (4) an ilium with larger supra-acetabular crest and proportionally shorter posterior blade; and (5) proximal tarsals that are more flared." de Ricqlès et al. (2003) and Padian et al. (2004) reported on its histology. Irmis (2005) stated "It is likely, given the morphological and temporal differences between the Petrified Forest and Ghost Ranch material, that they represent separate taxa (Parker and Irmis, in press). Therefore, we refer the Petrified Forest material to Coelophysis sp. pending description of the new material." Unfortunately, the resulting Parker and Irmis (2005) publication didn't comment on differences or taxonomy, still calling it Coelophysis sp.. Nesbitt et al. (2007) stated "Recent recovery of new specimens of the same taxon from equivalent strata in Petrified Forest National Park suggests that the Petrified Forest taxon is generally larger and more robust than the Ghost Ranch material" and referred it to Coelophysis sp. without justification. Spielmann et al. (2007) incorrectly stated "no specific differences have been enumerated in the literature" and provided high resolution photos of most pelvic and hindlimb elements. While they believed the morphological, temporal and geographical similarity with Coelophysis bauri and the Snyder Quarry coelophysid made it "most parsimonious to assign both to the same taxon" and thus referred UCMP 129618 to Coelophysis bauri, they only listed characters which would place it closer to Coelophysis than Dilophosaurus. Cabreira et al. (2016) first included the specimen in a phylogenetic analysis as the Petrified Forest theropod where it emerges in a trichotomy with Dilophosaurus and Liliensternus+Coelophysidae, but with a small sample size of coelophysoid-grade taxa and no averostrans this precise placement is poorly supported. Wang et al. (2017) included this in an independent analysis and recovered it as a coelophysoid just outside Coelophysidae. Ezcurra (2017) called this Padian's Coelophysis, and stated it differs from C. bauri (USNM 529376) in lacking a femoral extensor fossa. Marsh and Parker (2020) figured much of the specimen and referred it "to the Coelophysidae owing to the presence of the infrapopliteal ridge on the distal end of the femur and the coossifed proximal tarsals" although they mentioned these also occur in some ceratosaurs. Preliminary comparisons suggest it is a coelophysid based on the fused metatarsal III and distal tarsal III, but is not a coelophysine based on the rounded anteromedial corner of the femur in distal view.
Two of "several theropods" mentioned by Stocker et al. (2004) in an abstract, Parker and Irmis (2005) comment on "the partial skeletons of three coelophysoids. These specimens have not yet been completely prepared, though preliminary research has shown them to be identical to UCMP 129618." Discovered in 2004 and apparently not catalogued yet, Parker and Irmis figure the proximal femur of what would be PEFO 22983 and what is presumably the distal left femur of PEFO 33981 as "Coelophysis sp. ... from PFV 231." Nesbitt et al. (2007) mention two partial skeletons from the Giving Site (PEFO 33981 and 33983) as being the same taxon as UCMP 129618 based on greater size and robusticity than the C. bauri material from the Coelophysis Quarry. Marsh and Parker (2020) figured some of their elements and referred both to Coelophysidae for the same reasons as UCMP 129618. Parker and Irmis' third partial skeleton ended up being shuvosaurid PEFO 33953 (Parker, pers. comm. 2021). Most recently, Ezcurra et al. (2021) finds "PEFO 21373/UCMP 129618 and PEFO 33983 possess a unique combination of character states that allows assigning both to a single, new species of neotheropod. The third specimen [PEFO 33981] has a congruent morphology, but lacks preservation of key features, preventing an unambiguous referral to the new taxon. A phylogenetic analysis [presumably Nesbitt's modified by Ezcurra] found the new taxon as one of the earliest branching coelophysoids, outside Coelophysidae (Coelophysis + Megapnosaurus + "Syntarsus")."
Hunt and Wright (1999) note a specimen found in 1999 at the Jeremiah's Perch locality which they assign to the same species as UCMP 129618, stating it "is currently only partially excavated, but it includes at least femora, tibiae, cervical vertebrae, teeth as well as many other elements." Marsh and Parker (2020) stated it "was recently returned and prepared at PEFO."
References- Padian, 1986. On the type material of Coelophysis Cope (Saurischia: Theropoda) and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation). In Padian (ed.). The Beginning of the Age of Dinosaurs: Faunal Change Across the Triassic-Jurassic Boundary. Cambridge University Press. 45-60.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic dinosaur from New Mexico (USA). Paläontologische Zeitschrift. 65, 191-198.
Hunt, Santucci and McClelland, 1998. Preliminary results of the Dawn of the Dinosaurs Project at Petrified Forest National Park, Arizona. National Park Service Paleontological Research, Geological Resources Division Technical Report NPS/NRGRD/GRDTR-98/01. 135-137.
de Ricqlès, Padian and Horner, 2003. On the bone histology of some Triassic pseudosuchian archosaurs and related taxa. Annales de Paläontologie. 89(2), 67-101.
Padian, Horner and De Ricqlès, 2004. Growth in small dinosaurs and pterosaurs: The evolution of archosaurian growth strategies. Journal of Vertebrate Paleontology. 24(3), 555-571.
Stocker, Parker, Irmis and Shuman, 2004. New discoveries from the Upper Triassic Chinle Formation as the result of the ongoing paleontological inventory of Petrified Forest National Park. Journal of Vertebrate Paleontology, Program and Abstracts. 127.
Irmis, 2005. The vertebrate fauna of the Upper Triassic Chinle Formation in northern Arizona. In Nesbitt, Parker and Irmis (eds.). Guidebook to the Triassic Formations of the Colorado Plateau in Northern Arizona: Geology, Paleontology, and History. Mesa Southwest Museum, Bulletin. 9, 63-88.
Parker and Irmis, 2005. Advances in Late Triassic vertebrate paleontology based on new material from Petrified Forest National Park, Arizona. In Heckert and Lucas (eds.). Vertebrate Paleontology in Arizona. New Mexico Museum of Natural History and Science Bulletin. 29, 45-58.
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.
Cabreira, Kellner, Dias-da-Silva, da Silva, Bronzati, de Almeida Marsola, Müller, de Souza Bittencourt, Batista, Raugust and Carrilho, 2016. A unique Late Triassic dinosauromorph assemblage reveals dinosaur ancestral anatomy and diet. Current Biology. 26(22), 3090-3095.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
Ezcurra, 2017. A new early coelophysoid neotheropod from the Late Triassic of northwestern Argentina. Ameghiniana. 54, 506-538.
Marsh and Parker, 2020. New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils. PaleoBios. 37, 1-56.
Ezcurra, Marsh, Irmis and Nesbitt, 2021. A revision of coelophysoid theropod specimens from Petrified Forest National Park, Arizona (U.S.A.), reveals a new species from the Upper Triassic Chinle Formation. 34 Jornadas Argentinas de Paleontologia de Vertebrados, Libro de Resumenes. R16.

Shake-N-Bake coelophysid (Tykoski, 1997)
Sinemurian-Pliensbachian, Early Jurassic
Shake-N-Bake MCZ 40/78a, Silty Facies Member of the 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 "Styntarsus" kayentakatae by Rowe (1989). Initially catalogued 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. It is assigned to Coelophysidae here based on tibiotarsal fusion.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9(2), 125-136.
Tykoski, 1997. A new ceratosaurid theropod from the Early Jurassic Kayenta Formation of northern Arizona. Journal of Vertebrate Paleontology. 17(3), 81A-82A
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications for ceratosaurid phylogeny. Masters thesis, University of Texas at Austin. 217 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD thesis, University of Texas at Austin. 553 pp.

unnamed coelophysid (Simmons, 1965)
Sinemurian, Early Jurassic
Ta Ti, Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Material- (FMNH CUP 2089) vertebra?, distal humerus, proximal radius, proximal ulna, metacarpal fragments, manual phalangeal fragments
....(FMNH CUP 2090) partial tarsometatarsus
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 this is resolved as a coelophysid character here.
References- Simmons, 1965. The non-therapsid reptiles of the Lufeng Basin, Yunnan, China. Fieldiana. 15, 1-93.
Irmis, 2004. First report of Megapnosaurus (Theropoda: Coelophysoidea) from China. PaleoBios. 24(3), 11-18.

undescribed coelophysid (Jenny, Jenny-Deshusses, Le Marrec and Taquet, 1980)
Toarcian, Early Jurassic
Wazzant Formation, Wazzant, Morocco
Material- (MNHN coll.) (~1.5 m) cervical vertebra, dorsal vertebrae, sacral vertebrae, caudal vertebrae, pelvis, hindlimbs including femur, tibia, fibula, partial astragalus, calcaneum, distal tarsal IV, metatarsal I, phalanx I-1, ungual I, metatarsal II (~114 mm), incomplete phalanx II-1, metatarsal III (~111 mm) fused with distal tarsal III, phalanx III-1 (~24 mm), proximal phalanx III-2, metatarsal IV (~105 mm), metatarsal V, pedal phalanx (~19 mm), phalanx (~17 mm), pedal ungual (~18 mm)
Comments- Discovered in 1978 (Taquet, 1986), Jenny et al. (1980) gave a brief initial description of this "small Theropod skeleton (about 1.50 m in length)" (translated), noting "at least the posterior half of the skeleton is present with the caudal, sacral, dorsal vertebrae, the pelvis and the two hind legs." They concluded "If the distal end of the femur and the proximal end of the tibia closely resemble (but are five times smaller) the corresponding ends of the femur and tibia of Allosaurus (=Antrodemus) from the Upper Jurassic of North America, the astragalus is different and the metatarsals are very slender. This is why it seems to us premature in the current state of our study to place this theropod among the carnosaurians rather than among the coelurosaurians." Taquet (1985) later figured the pes as "Liassic coelurosaur from Morocco", noting "The posterior view enables us to observe the elements of the first digit in articulation with practically no displacement." It should be noted both of these studies used 'coelurosaur' in the Hueneian sense which included coelophysoids. Taquet (1986) stated "The preparation of this specimen is in progress. The priority given to the preparation and casting of the Cetiosaurus of Wawmda [= Atlasaurus type] for the Museum of Earth Sciences in Rabat did not allow us to carry out the clearing work with the desired speed" (translated). Allain et al. (2007) state it "is currently under preparation in the Muséum National d’Histoire Naturelle, in Paris. It differs clearly from Berberosaurus in its smaller size, the longer cervical vertebral centra, and the absence of a sulcus on the medial surface of the fibula." Benson (2010) noted based on Taquet's figure it "appears to be a non-tetanuran theropod" based on the "low, block-like ascending process of the astragalus." This specimen is provisionally assigned to Coelophysidae based on the apparent fusion between distal tarsal III and metatarsal III.
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, 1985. 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.
Taquet, 1986. Découvertes récentes de Dinosaures au Maroc. In Taquet and Sudre (eds.). Les Dinosaures de la Chine à la France. Musée d'Histoire Naturelle de Toulouse. 39-43.
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.
Benson, 2010. The osteology of Magnosaurus nethercombensis (Dinosauria, Theropoda) from the Bajocian (Middle Jurassic) of the United Kingdom and a re-examination of the oldest records of tetanurans. Journal of Systematic Palaeontology. 8(1), 131-146.

Lucianovenator Martinez and Apaldetti, 2017
= "Lucianosaurus" Martinez and Apaldetti, 2017 online preoccupied Hunt and Lucas, 1994
L. bonoi Martinez and Apaldetti, 2017
= "Lucianosaurus bonoi" Martinez and Apaldetti, 2017 online
Late Norian-Rhaetian, Late Triassic
Quebrada del Barro Formation, San Juan, Argentina
Holotype- (PVSJ 906) distal axial rib, partial third cervical vertebra, distal third cervical rib, fourth cervical vertebra (49.8 mm) fused to cervical rib, fifth cervical vertebra (54.7 mm) fused to cervical rib, sixth cervical vertebra (57 mm) fused to cervical rib, seventh cervical vertebra (56.6 mm) fused to cervical rib, eighth cervical vertebra (54 mm), ninth cervical vertebra (52.8 mm), tenth cervical vertebra (40.8 mm), first dorsal vertebra (38 mm), second dorsal vertebra (31.6 mm), incomplete third dorsal vertebra (32 mm), fourth dorsal vertebra (31.3 mm), thirteenth dorsal vertebra (37.1 mm), proximal first to third dorsal ribs, synsacrum (154 mm; 36.2, 28.6, 26.3, 30.2, 32 mm), first caudal vertebra (32.6 mm), incomplete ilia (145.6 mm) fused with proximal ischium
Paratypes- (PVSJ 899) incomplete synsacrum (~107 mm; ?, 25.9, 22.2, 21.7, 25.9 mm), first caudal vertebra (24.8 mm), incomplete ilia fused with proximal pubes and proximal ischia
(PVSJ 1013) partial synsacrum (~908 mm; ?, 18.5, 16.3, 20.5, 21.7 mm)
(PVSJ 1084) synsacrum (129 mm; ?, ?, ?, 26.2, 25.3 mm), first caudal vertebra, incomplete ilia fused with proximal ischia
Referred- ?(PVSJ 1004) proximal tibia (Martinez and Apaldetti, 2017)
Diagnosis- (after Martinez and Apaldetti, 2017) cervical neural arches with series of three deep fossae, two blind pockets located within the prezygapophyseal centrodiapophyseal fossa and a third, communicating with the internal cavity of the neural arch, located within the centropostzygapophyseal fossa; anteroposteriorly elongated fossa on the edge of the posterior centrodiapophyseal lamina that progressively increases in depth and size from C3 to C9; length of the anterior cervical ribs equivalent to five cervical centra.
Comments- This material was discovered after 2000 and initially described as Lucianosaurus bonoi (Martinez and Apaldetti, 2017 online) in an April 17 2017 preprint, which Creisler (DML, 2017) noted was preoccupied by the archosauromorph tooth taxon Lucianosaurus wildi (Hunt and Lucas, 1994). Martinez and Apaldetti (2017) quickly corrected their draft on May 11 2017 with the new genus name Lucianovenator. Lucianosaurus was never registered with ZooBank, though Lucianovenator has a ZooBank entry but no explicit statement of ZooBank registration in its publication. Thus both names were considered nomina nuda pending physical publication (ICZN Article 8.5.3), which occured in November 2017 for Lucianovenator.
Martinez and Apaldetti recover Lucianovenator as a coelophysid in a trichotomy with Camposaurus and Coelophysis? rhodesiensis using a version of Nesbitt et al.'s dinosaur analysis. Only one or two steps are needed to shift the genus to another position within Coelophysidae, but moving it outside the clade requires 8 or more steps.
References- Hunt and Lucas, 1994. Ornithischian dinosaurs from the Upper Triassic of the United States. In Fraser and Sues (eds.). In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods. Cambridge University Press. 227-241.
Creisler, DML 2017. https://web.archive.org/web/20191030050839/http://dml.cmnh.org/2017Apr/msg00094.html
Martinez and Apaldetti, 2017 online. A Late Norian-Rhaetian coelophysid neotheropod (Dinosauria, Saurischia) from the Quebrada del Barro Formation, northwestern Argentina. Ameghiniana. (advance online publication)
Martinez and Apaldetti, 2017. A Late Norian-Rhaetian coelophysid neotheropod (Dinosauria, Saurischia) from the Quebrada del Barro Formation, northwestern Argentina. Ameghiniana. 54(5), 488-505.

"Syntarsus" kayentakatae Rowe, 1989
= Coelophysis kayentakatae (Rowe, 1989) Bristowe and Rowe, 2004
= Megapnosaurus kayentakatae (Rowe, 1989) Tykoski and Rowe, 2004
Sinemurian-Pliensbachian, Early Jurassic
Rock Head North MNA 555-3, Silty Facies Member of the Kayenta Formation, Arizona, US
Holotype- (MNA.V.2623 in part; field number 18/78a) (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), semilunate carpal, 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)
Paratype- (MNA.V.2623 in part) (15-20% larger than holotype; at least two robust individuals) 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
Sinemurian-Pliensbachian, Early Jurassic
Paiute Canyon General TMM 43648, Silty Facies Member of the Kayenta Formation, Arizona, US
Referred- (TMM 43648-9) (adult) material including distal tibia (Tykoski, 2005)
Sinemurian-Pliensbachian, Early Jurassic
Gold Spring Wash TMM 43669, Silty Facies Member of the Kayenta Formation, Arizona, US
(TMM 43669-3) (young subadult) tibia, astragalocalcaneum (~23 mm trans) (Tykoski, 2005)
Sinemurian-Pliensbachian, Early Jurassic
TMM 43688, Silty Facies Member of the Kayenta Formation, Arizona, US
(TMM 43688-1; field number TR 97/12) (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 (Tykoski, 1998)
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; MCZ 8817) 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. Barta et al. (2018) suggested metacarpal I and phalanx II-2 of Tykoski may be phalanx I-1 or II-1 and II-1 or III-2/3 respectively.
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. Excurra (2012) did find the two to be sister taxa and recommended synonymy based on an unpublished analysis though.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9(2), 125-136.
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications for ceratosaurid phylogeny. 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, 2001 online. Syntarsus kayentakatae, Digital Morphology. http://digimorph.org/specimens/Syntarsus_kayentakatae/
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 (eds.). The Dinosauria Second Edition. University of California Press. 47-70.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD thesis. University of Texas at Austin. 553 pp.
Gay, 2010. Notes on early Mesozoic theropods. Lulu Press. 44 pp.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Barta, Nesbitt and Norell, 2018 (online 2017). The evolution of the manus of early theropod dinosaurs is characterized by high inter- and intraspecific variation. Journal of Anatomy. 232(1), 80-104.

"Syntarsus" "mexicanum" Hernandez, 2002
Toarcian, Early Jurassic
Casa de Fidencio, 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
Toarcian, Early Jurassic
Rene's Skull, La Boca Formation, Mexico
?(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 "Syntarsus" based on the antitrochanter projecting into the acetabulum (though this is also present in Coelophysis bauri and Liliensternus). It differs in having ilia contact sacral neural spines two and three. The skull fragments may not belong to the same taxon and were thought to be a neoceratosaur or maniraptoran by Munter due to their pneumatized paroccipital roots. However, this has since been found to exist in coelophysids such as "S." kayentakatae. 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 (excluded due to lacking fused sacral neural spines), and Rauhut's (2003) matrix where it was in the clade of Coelophysis (the latter including 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. Ezcurra (2012) found the specimen to be closer to Coelophysis, Segisaurus and Camposaurus than kayentakatae based on a large unpublished analysis.
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.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.

Procompsognathinae Nopcsa, 1923
Definition- (Procompsognathus triassicus <- Coelophysis bauri) (Sereno, online 2005; 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 coelophysine Coelophysis. This was not found in future studies (e.g. Tykoski, 2005; Ezcurra and Novas, 2007; Ezcurra, 2012), 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. Journal of Vertebrate Paleontology. 24(3), 117-118.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra and Novas, 2007 (online 2006). Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.

Panguraptor You, Azuma, Wang, Wang and Dong, 2014
P. lufengensis You, Azuma, Wang, Wang and Dong, 2014
Hettangian, Early Jurassic
Shawan Member (Dull Purplish Beds) of the Lufeng Formation, Yunnan, China
Material- (LFGT-0103) (~2 m; subadult) incomplete skull (~124 mm), incomplete mandible, atlas, axis (18.5 mm), third cervical vertebra (24 mm), fourth cervical vertebra (28 mm), fifth cervical vertebra (29 mm), sixth cervical vertebra (31 mm), seventh cervical vertebra (30.5 mm), eighth cervical vertebra (28.5 mm), ninth cervical vertebra (28 mm), tenth cervical vertebra (23 mm), cervical ribs, first dorsal neural spine, second dorsal neural spine, second dorsal neural spine, third dorsal neural spine, fourth dorsal neural spine, fifth dorsal neural spine, sixth dorsal neural arch, seventh dorsal neural arch, eighth dorsal neural arch, ninth dorsal vertebra (24 mm), tenth dorsal vertebra (26 mm), eleventh dorsal vertebra (26 mm), twelfth dorsal vertebra (25 mm), thirteenth dorsal vertebra (23 mm), over eleven partial dorsal ribs, first sacral vertebra, incomplete scapula (86 mm), distal humerus, radius (~57 mm), ulna, metacarpal I, phalanx I-1 (17 mm), manual ungual I (21 mm), metacarpal II (36 mm), phalanx II-1 (18 mm), phalanx II-2 (22 mm), manual ungual II, metacarpal III (34 mm), phalanx III-1 (10 mm), phalanx III-2 (8 mm), phalanx III-3 (9 mm), metacarpal IV (22 mm), phalanx IV-1, phalanx IV-2(?), partial ilium, distal ischia, femora (one partial; 164 mm), tibia (182 mm), fibula (178 mm), astragalus, calcaneum, distal tarsal IV, metatarsal III, phalanx III-1, phalanx III-2 (22 mm), phalanx III-3 (23 mm), pedal ungual III (17 mm), metatarsal IV (98 mm), phalanx IV-1 (18 mm), phalanx IV-2 (17 mm), phalanx IV-3 (13.5 mm), metatarsal V (41 mm)
Diagnosis- (after You et al., 2013) large antorbital fenestra; anterodorsal-posteroventral ridge on lateral surface of maxilla, within antorbital fossa; elliptical, laterally facing fenestra posterodorsal to diagonal ridge (also in Zupaysaurus); promaxillary fenestra; long maxillary body; hooked anteromedial corner of distal tarsal IV.
Comments- Panguraptor was discovered in 2010, initially described in an abstract by You et al. (2013), then fully described and named by You et al. (2014).
You et al. (2013) found this taxon to be more closely related to Coelophysis bauri than C. rhodesiensis, while in the official description You et al. (2014) found it to be outside the bauri-rhodesiensis clade, but closer to them than kayentakatae.
References- You, Azuma, Wang and Dong, 2013. A new coelophysoid theropod dinosaur from the Early Jurassic Lufeng Formation og Yunnan Province, China. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 242.
You, Azuma, Wang, Wang and Dong, 2014. The first well-preserved coelophysoid theropod dinosaur from Asia. Zootaxa. 3873(3), 233-249.

Procompsognathus Fraas, 1913
P. triassicus Fraas, 1913
= Hallopus "celerrimus" Fraas, 1912
Middle Norian, Late Triassic
Pfaffenhofen, Middle Löwenstein Formation, Germany
Holotype- (SMNS 12591) (.96 m) (robust) partial cervical vertebrae 5-10 (~14-16 mm), 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, radius, ulna (34.2 mm), radiale?, metacarpal I (~4.8 mm), phalanx I-1 (~5.6 mm), manual ungual I, metacarpal II (~11 mm), partial phalanx II-1 (~6.5 mm), incomplete metacarpal III, partial metacarpal IV, partial ilium, pubes, 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)
?...(SMNS 12591a) incomplete skull
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.
Procompsognathus was found in 1909. It was described based on a postcranial skeleton (SMNS 12591) and skull (SMNS 12591a), which were long assumed to belong to the same individual but probably do not based on the differently colored matrix and small size of the skull compared to Coelophysis' proportions (Knoll, 2008). Huene (1921) referred another skull (SMNS 12352) and manus (SMNS 12352a) to the taxon, but these are now agreed to be crocodylomorphan (Welles, 1984; Sereno and Wild, 1992; Knoll and Rohrberg, 2012). Huene (1921) also referred parts of the Saltoposuchus type slab (SMNS 12597) to Procompsognathus, but these were agreed to be Saltoposuchus by Crush (1984).
Sereno and Wild (1992) considered the skull SMNS 12591a to belong to Saltoposuchus connectens, while Chatterjee (1993, 1998) believed it to be theropod. 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. Knoll and Schoch (2006) tentatively stated the skull was tetanurine based on a supposed maxillary fenestra, yet Knoll and Rohrberg (2012) figure it as Theropoda indet.. Ezcurra (2012) found Procompsognathus to be a non-coelophysid coelophysoid sensu stricto in a large unpublished analysis, though it was unreported if the skull was coded.
References- Fraas, 1912. Die schwäbischen Dinosaurier. Jahreshefte des Vereins für Vaterländische Naturkunde in Württemberg. 68, 66-67.
Fraas, 1913. Die neuesten Dinosaurierfunde in der schwabischen Trias. Naturwissenschaften. 45, 1097-1100.
Huene, 1921. Neue Pseudosuchier und Coelurosaurier aus dem württembergischen Keuper. Acta Zoologica. 2, 329-403.
Walker, 1961. Triassic reptiles from the Elgin area: Stagonolepis, Dasygnathus and their allies. Philosophical Transactions of the Royal Society of London, Series B. 244, 103-204.
Ostrom, 1981. Procompsognathus - theropod or thecodont? Palaeontographica, Abteilung A. 175, 179-195.
Crush, 1984. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology. 27, 131-157.
Sereno and Wild, 1992. Procompsognathus: Theropod, "thecodont" or both? Journal of Vertebrate Paleontology. 12(4), 435-458.
Chatterjee, 1993. Procompsognathus from the Triassic of Germany is not a crocodylomorph. Journal of Vertebrate Paleontology. 13(3), 29A.
Chatterjee, 1998. Reassessment of Procompsognathus skull. In Wolberg, Gittis, Miller, Carey and Raynor (eds.). Dinofest International. 6.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD dissertation. University of Bristol. 440 pp.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. Journal of Vertebrate Paleontology. 24(3), 117A-118A.
Knoll and Schoch, 2006. Does Procompsognathus have a head? Systematics of an enigmatic Triassic taxon. Journal of Vertebrate Paleontology. 26(3), 86A.
Knoll, 2008. On the Procompsognathus postcranium (Late Triassic, Germany). Geobios. 41(6), 779-786.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Knoll and Rohrberg, 2012. CT scanning, rapid prototyping and re-examination of a partial skull of a basal crocodylomorph from the Late Triassic of Germany. Swiss Journal of Geosciences. 105(1), 109-115.

Coelophysinae Nopcsa, 1928
Definition- (Coelophysis bauri <- Procompsognathus triassicus) (Ezcurra, 2017; modified from Sereno, 1998)
= Segisaurinae Camp, 1936 vide Kalandadze and Rautian, 1991
= Syntarsiinae Kalandadze and Rautian, 1991
= Coelophysidae sensu Tykoski and Rowe, 2004
Definition- (Coelophysis bauri + Coelophysis rhodesiensis) (modified)
Diagnosis- (suggested) promaxillary fenestra absent; lateral dentary groove absent; humerus straight in lateral view; angled anteromedial corner of femur in distal view.
Comments- Although 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 and Elaphrosaurus, to separate them from the halticosaurine Liliensternus and Dilophosaurus. Sereno (1998) then used it to group Coelophysis bauri and Megapnosaurus rhodesiensis, 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 rhodesiensis + bauri clade, excluding Liliensternus. Taking these prior uses into account, Coelophysinae is here used for the rhodesiensis + bauri 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, 2007) and is equivalent to Sereno's Coelophysinae in content. Ezcurra (2012) found this to only include Coelophysis bauri however, with kayentakatae more basal, and Camposaurus and rhodesiensis as segisaurines.
Segisaurinae was based on Camp's family Segisauridae and is used here for the clade formed by Segisaurus, "Syntarsus" kayentakatae and possibly Camposaurus recovered by Tykoski (2005). Such a clade was not recovered by Ezcurra and Novas (2006), where "Syntarsus" kayentakatae was instead closer to Coelophysis than to Segisaurus. Similarly, it was not recovered in Ezcurra (2012) where Camposaurus and rhodesiensis are segisaurines, and kayentakatae is a basal coelophysid.
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.
Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of Arizona. University of California Publications in Geological Sciences. 24(2), 39-56.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 464 pp.
Kalandadze and Rautian, 1991. Late Triassic zoogeography and reconstruction of the terrestrial tetrapod fauna of North Africa. Paleontological Journal. 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 Thesis. University of Texas at Austin. 553 pp.
Ezcurra and Novas, 2007 (online 2006). Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Ezcurra. 2017. A new early coelophysoid neotheropod from the Late Triassic of northwestern Argentina. Ameghiniana. 54, 506-538.

unnamed coelophysine (Morales, 1994)
Early Hettangian, Early Jurassic
Thybony Site MNA 854-1, Dinosaur Canyon Member of the Moenave Formation, Arizona, US
Material- (MNA.V.1968) teeth(?), dorsal ribs, gastralia, incomplete ilium, proximal pubis, proximal ischium, distal ischia, proximal femur, phalanges(?)
(MNA.V.2588) (centra 34-37 mm) fourth sacral centrum, fifth sacral centrum, first caudal centrum, second caudal centrum, third caudal centrum, ilium (196 mm), proximal pubis, ischia (172 mm)
Comments- Discovered in 1986, this material was first reported in an abstract by Morales (1994) who stated it "may represent two individuals" and mentioned "phalanges; and some isolated teeth" not noted in the eventual description by Lucas and Heckert (2001) but which may be some of "the remaining bones" of MNA.V.1968 that are said to be undiagnostic. When Morales stated "Distally the pubes are fused forming a rod-like (not plate-like) pubic shaft and ending as a single united expansion (a knob, not a pubic "foot")", he was describing the distal ischia of MNA.V.1968. Also, Morales' statement "the part of the pubic plate where the obturator foramen and pubic fenestra might lie, is not preserved" is incorrect, as Lucas and Heckert find "One foramen is well demarcated, but whether this is the obtuator foramen or an additional pubic fenestra is uncertain."
Morales (1994) believed "The shape and relative size of the brevis shelf of the ilium and the degree of ventrolateral expansion of the supracetabular crest is more similar to Syntarsus" than Coelophysis, but did not assign it to a genus. Lucas and Heckert (2001) thought the level of the ilioischial suture was "different from the photographic illustration of Coelophysis by COLBERT (1989: fig. 76)", but must have mistaken a more ventral break in that figure for the suture, which is not visible and thus probably completely fused. Similarly, when they state the inturned femoral head is "more marked than that of Coelophysis illustrated by COLBERT (1989: fig. 80)", the latter is an innacurate drawing as detailed by Downs. They conclude "the pelves described here closely conform to descriptions of Syntarsus rhodesiensis and S. kayentakatae (RAATH 1977, TYKOSKI 1998), so we assign them to Syntarsus sp.", also noting they are larger than the Shake-N-Bake coelophysid. Lucas and Heckert also state "the pubes fuse to the ischia slightly posterior of the midpoint of the acetabulum, which is positioned farther posteriorly than in Syntarsus or Coelophysisand may represent a unique or species-level distinction." Wang et al. (2017) is the only publication to include the specimen in an analysis (as "Moenave Coelophysoid"), where it emerged as a coelophysid in a polytomy with Camposaurus, Megapnosaurus and kayentakatae. It is assigned to Coelophysinae here based on the shallow ilium with concave dorsal margin.
References- Morales, 1994. First dinosaur body fossils from the Lower Jurassic Dinosaur Canyon Member, Moenave Formation of northeastern Arizona. Journal of Vertebrate Paleontology. 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.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.

Camposaurus Hunt, Lucas, Heckert, Sullivan and Lockley, 1998
C. arizonensis Hunt, Lucas, Heckert, Sullivan and Lockley, 1998
Middle Norian, Late Triassic
Placerias Quarry UCMP A269, Sonsela Member of the Chinle Formation, Arizona, US
Holotype- (UCMP 34498) distal tibiotarsi fused with distal fibulae
Diagnosis- (after Hunt et al., 1998) ventral margin incipiently concave in anterior view (also in GR 227 and NMMNH P-29168).
(after Ezcurra and Brusatte, 2011) posterior edge of tibial articular surface for fibula prominent and takes the form of a sharp longitudial ridge (also in GR 227; unreported in Coelophysis bauri); strongly developed, anteriorly bowed, diagonal tuberosity on tibial medial surface (also in GR 227; poorly developed in rhodesiensis); astragalus without strong anterior projection of medial condyle (also in GR 227 and NMMNH P-29168).
Other diagnoses- Hunt et al. (1998) also listed "femoral head more rectangular and dorsal centra more waisted in ventral view" based on paratype specimens UCMP 139622, MNA.V.2777 and UCMP 177314, but these are not referred to Camposaurus here.
Comments- The holotype tibiotarsi were collected in 1934 and first published by Murry and Long (1989), who said "Theropod bones (tarsi, sacral and distal vertebrae) are present in the University of California Museum of Paleontology collections from the Placerias Quarry." Lucas et al. (1992) later figured and mentioned them as "fused ankles that consist of the distal ends of co-ossified tibiae-fibulae-astragalae. These bones probably pertain to ceratosaurian theropods." Long and Murry (1995) figured one of the tibiotarsi as Ceratosauria indet. and described them as Theropoda indet., but noted the straight distal margin in anterior/posterior view "differs from that seen within other primitive theropods" including UCMP 129618, and that the less concave anterior margin later found to be diagnostic was different from Liliensternus and Dilophosaurus at least. Hunt et al. (1998) included UCMP 34498 in "A large number of dissociated, similar-sized theropod postcranial specimens, with no duplication of elements" that they believed "represent a single individual" of their new taxon Camposaurus arizonensis (see below for paratype material). They referred Camposaurus to Ceratosauria sensu lato based on the fused astragalocalcanea, and thought it was most similar to Coelophysis and rhodesiensis (their Rioarribasaurus and Syntarsus) in the fused tibiotarsus. Downs (2000) states Camposaurus "was founded on hind limb and tarsal characters which are contrasted with Coelophysis characters illustrated [incorrectly by L. Darling] in Colbert's monograph. Camposaurus appears to fall easily within the range of variation seen in the Ghost Ranch sample of Coelophysis and is thus a nomen dubium." Irmis (2005) agreed, stating "direct comparison of Camposaurus with casts of corresponding elements of Coelophysis bauri from the Ghost Ranch Coelophysis Quarry shows that they are identical." Nesbitt et al. (2005, 2007) agreed it cannot be distinguished based on the straight distal margin, claiming they "could not corroborate this difference when directly comparing 'Camposaurus' to Coelophysis bauri (AMNH FR 30614 and AMNH FR 30615)", but their own figure 5 shows the opposite. Similarly, they say "One difference between 'Camposaurus,' some specimens of Coelophysis bauri and all other theropods is that in ventral view, the concave depression on the anterior side of the astragalus is much stronger and more abrupt in Coelophysis bauri; however, some specimens of Coelophysis bauri have a morphology identical to 'Camposaurus'", referring to AMNH 30614 in their Figure 5. Yet that specimen clearly has the medially extensive anterior groove of Coelophysis bauri and most other theropods, and is merely compressed anteroposteriorly to make the depression appear shallow. Tykoski (2005) stated the presence of fibulocalcanear fusion is otherwise only seen in "Syntarsus" kayentakatae, though otherwise the material could not be placed more precisely within the Coelophysis<Liliensternus clade. The holotype was reanalyzed by Ezcurra and Brusatte (2011), who argued several characters can distinguish Camposaurus from other named taxa, including the straighter distal and anterior astragalocalcanear margins. The authors added Camposaurus to Nesbitt's dinosauromorph analysis, finding it to be a coelophysid closer to rhodesiensis than to Coelophysis bauri and kayentakatae. Similarly, Ezcurra (2012) found Camposaurus to be closer to rhodesiensis and Segisaurus than to Coelophysis bauri and kayentakatae based on a large unpublished analysis. Wang et al. (2017) recovered it as a coelophysid in a polytomy with rhodesiensis and kayentakatae.
Referred material- Hunt et al. (1998) included several elements from the same locality as paratypes, but future authors such as Nesbitt et al. (2007) and Ezcurra and Brusatte (2011) have excluded these "Given that the holotype and referred specimens were discovered in a quarry containing the bones of many taxa and individuals, and that there is no clear association between them."
Proximal femur UCMP 139622 (UCMP uncatalogued in Hunt et al., 1998) has since been assigned to Saurischia by Irmis (2005) and the more rectangular head suggested by Hunt et al. as diagnostic is indeed different from coelophysoid-grade taxa. Dorsal centra MNA.V.3091 ('MNA V2777' in Hunt et al.) and UCMP 177314 (UCMP uncatalogued in Hunt et al.), partial synsacrum UCMP 138591, and fragmentary synsacra UCMP 178047, 178048 and 178049 (all UCMP uncatalogued in Hunt et al.) were all placed in Archosauria indet. by Nesbitt et al. (2007) as "they are equally comparable to many dinosauriforms as well as Shuvosaurus."
Proximal pubis UCMP 177318 was tentatively referred to Camposaurus by Hunt et al. (1998), but is reassigned to Archosauria indet. here.
Tykoski (2005) referred to "a piece of the right pelvic girdle that includes the acetabular border formed by fusion of the pubic peduncle of the ilium and the proximal pubis (UCMP 25791)" as possibly from the Camposaurus type individual. This had only been previously published as a Rutiodon specimen preserving a "Pubic tubercle reduced to a rugosity" by Hutchinson (2001), and is in the UCMP online catalogue as a reptilian ulna discovered in 1934. It's possible Tykoski confused this with UCMP 177318 noted above, but this is a left proximal pubis that shows no evidence of fusion with the ilium.
Nesbitt et al. (2007) say distal femur UCMP 25834 and distal tibia UCMP 25820 from the Placerias Quarry may belong to Camposaurus, but neither has been figured or supported with autapomorphies, although the tibia could potentially be examined for Camposaurus' distinctive ridge and tuberosity.
Ezcurra and Brusatte (2011) noted Hayden Quarry coelophysid tibiotarsus GR 227 shares all of their proposed Camposaurus apomorphies, which could mean the Hayden Quarry material is referrable to that taxon. This is not formalized here as the Hayden material is undescribed and from a higher stratigraphic level than Camposaurus.
References- 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. In Lucas and Hunt (eds.). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History. 29-64.
Lucas, Hunt and Long, 1992. The oldest dinosaurs. Naturwissenschaften. 79(4), 171-172.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the southwestern Unites States. New Mexico Museum of Natural History and Science Bulletin. 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. New Mexico Museum of Natural History and Science Bulletin. 17, 33-37.
Hutchinson, 2001. The evolution of pelvic osteology and soft tissues on the line to extant birds (Neornithes). Zoological Journal of the Linnean Society. 131, 123-168.
Irmis, 2005. The vertebrate fauna of the Upper Triassic Chinle Formation in northern Arizona. In Nesbitt, Parker and Irmis (eds.). Guidebook to the Triassic Formations of the Colorado Plateau in Northern Arizona: Geology, Paleontology, and History. Mesa Southwest Museum, Bulletin. 9, 63-88.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. Journal of Vertebrate Paleontology. 25(3), 96A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD thesis. 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.
Ezcurra and Brusatte, 2011. Taxonomic and phylogenetic reassessment of the early neotheropod dinosaur Camposaurus arizonensis from the Late Triassic of North America. Palaeontology. 54(4), 763-772.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.

cf. Camposaurus (Irmis, Nesbitt and Downs, 2006)
Late Norian, Late Triassic
Hayden Quarry, Petrified Forest Member of the Chinle Formation, New Mexico, US
(GR 211) astragalocalcaneum (Nesbitt and Stocker, 2008)
(GR 227) tibiotarsus, fibula (Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007)
(GR 1033) semilunate carpal, complete manus including metacarpal I, metacarpal II and metacarpal III (Barta, Nesbitt and Norell, 2018)
(GR coll.) (several individuals) (Irmis, Nesbitt and Downs, 2006)
(H2-069-060602) ilium (Whiteside, Lindstrom, Irmis, Glasspool, Schaller, Dunlavey, Nesbitt, Smith and Turner, 2015)
Comments- Irmis et al. (2006) announced "several individuals of a large coelophysoid theropod" from the Hayden quarry, of which a tibiotarsus was figured by Irmis et al. (2007). Nesbitt and Stocker (2008) mention astragalocalcaneum GR 211, which was also listed in the appendix of Whiteside et al. (2015) as a coelophysid. Barta et al. (2018) figure a metacarpus GR 1033 that fuses distal carpal III to the semilunate carpal. The material has yet to be described. Ezcurra and Brusatte (2011) noted GR 227 shares all of their proposed Camposaurus apomorphies, which could mean the Hayden quarry material is referrable to that taxon. This is not formalized here as the Hayden material is undescribed and from a higher stratigraphic level than Camposaurus.
References- Irmis, Nesbitt and Downs, 2006. A new Upper Triassic vertebrate quarry from the Chinle Formation of northern New Mexico with a unique and exceptionally diverse tetrapod fauna. Journal of Vertebrate Paleontology. 26(3), 81A.
Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science. 317, 358-361.
Nesbitt and Stocker, 2008. The vertebrate assemblage of the Late Triassic Canjilon Quarry (northern New Mexico, USA), and the importance of apomorphy-based assemblage comparisons. Journal of Vertebrate Paleontology. 28(4), 1063-1072.
Ezcurra and Brusatte, 2011. Taxonomic and phylogenetic reassessment of the early neotheropod dinosaur Camposaurus arizonensis from the Late Triassic of North America. Palaeontology. 54(4), 763-772.
Whiteside, Lindstrom, Irmis, Glasspool, Schaller, Dunlavey, Nesbitt, Smith and Turner, 2015. Extreme ecosystem instability suppressed tropical dinosaur dominance for 30 million years. Proceedings of the National Academy of Sciences. 112(26), 7909-7913.
Barta, Nesbitt and Norell, 2018 (online 2017). The evolution of the manus of early theropod dinosaurs is characterized by high inter- and intraspecific variation. Journal of Anatomy. 232(1), 80-104.

cf. Coelophysis bauri (Heckert, Zeigler, Lucas, Rinehart and Harris, 2000)
Late Norian, Late Triassic
Snyder Quarry NMMNH L-3845, Petrified Forest Member of the Chinle Formation, New Mexico, US
(NMMNH P-29046) femora (155.5 mm), tibia (164 mm), proximal tibia, proximal fibula (Heckert, Zeigler, Lucas, Rinehart and Harris, 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, Zeigler, Lucas, Rinehart and Harris, 2000)
(NMMNH P-29168) partial tibiotarsus fused to fibula (Heckert, Zeigler, Lucas, Rinehart and Harris, 2000)
?(NMMNH P-30779) dorsal vertebra (Heckert, Zeigler, Lucas, Rinehart and Harris, 2000)
?(NMMNH P-30780) dorsal vertebra (Heckert, Zeigler, Lucas, Rinehart and Harris, 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, Zeigler, Lucas, Rinehart and Harris, 2000)
(NMMNH P-31293) cervical rib, incomplete tibia (Heckert, Zeigler, Lucas, Rinehart and Harris, 2000)
(NMMNH P-31661) cervical ribs, incomplete sacrum, partial scapulocoracoid, radius (51.8 mm), metacarpal, pedal ungual (Heckert, Zeigler, Lucas, Rinehart and Harris, 2000)
(NMMNH P-54617) tibia (Spielmann, Lucas, Rinehart, Hunt, Heckert and Sullivan, 2007)
(NMMNH P-54618) distal femur (Spielmann, Lucas, Rinehart, Hunt, Heckert and Sullivan, 2007)
(NMMNH P-54619) distal femur (Spielmann, Lucas, Rinehart, Hunt, Heckert and Sullivan, 2007)
(NMMNH P-54620) femur (245 mm) (Spielmann, Lucas, Rinehart, Hunt, Heckert and Sullivan, 2007)
Comments- 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 coelophysoids, and a square anterior end on the antorbital fossa as in Zupaysaurus and 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 Coelophysis rhodesiensis) 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. Wang et al. (2017) are the only publication to add the material to a phylogenetic analysis, recovering it as the most basal coelophysoid, outside Coelophysidae.
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.
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.
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.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.

Coelophysis Cope, 1889
= Syntarsus Raath, 1969 (preoccupied Fairmaire, 1869)
= Rioarribasaurus Hunt and Lucas, 1991
= Megapnosaurus Ivie, Slipinski and Wegrzynowicz, 2001
Comments- "Syntarsus"/Megapnosaurus 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, Ivie et al. discovered that Syntarsus was preoccupied by a zopherid beetle (Fairmaire, 1869). The entomologists who determined this attempted unsuccessfully to contact Raath so that he could rename it, and ended up renaming it themselves. 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, a large unpublished analysis by Ezcurra (2012) has found rhodesiensis to be a segisaurine, complicating taxonomy if this is found to be well supported.
References- Fairmaire, 1869. Notes sur les Coleopteres recueillis par Charles Coquerel a Madagascar et sur les cotes d'Afrique. 2e Partie. Annales de la Societe Entomologique de France, 4 Serie. 9, 179-260.
Cope, 1889. On a new genus of Triassic Dinosauria. American Naturalist. 23, 626.
Raath, 1969. A new coelurosaurian dinosaur from the Forest Sandstone of Rhodesia. Arnoldia. 4(28), 1-25.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic dinosaur from New Mexico (USA). Paläontologische Zeitschrift. 65, 191-198.
Downs, 2000. Coelophysis bauri and Syntarsus rhodesiensis compared, with comments on the perparation and preservation of fossils from the Ghost Ranch Coelophysis Quarry. New Mexico Museum of Natural History and Science Bulletin. 17, 33-37.
Ivie, Slipinski and Wegrzynowicz, 2001. Generic homonyms in the Colydiinae (Coleoptera: Zopheridae). Insecta Mundi. 15, 63-64.
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.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
C. bauri (Cope, 1887a) Cope, 1889
= Coelurus bauri Cope, 1887a
= Tanystropheus bauri (Cope, 1887a) Cope, 1887b
= Rioarribasaurus colberti Hunt and Lucas, 1991
= Syntarsus colberti (Hunt and Lucas, 1991) Paul, 1993
Early Rhaetian, Late Triassic
Coelophysis Quarry NMMNH L-3115, '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), metacarpal V, 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, radiale, ulnare, intermedium?, semilunate carpal, distal carpal III, manus including metacarpal I, metacarpal II and metacarpal III, 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, radiale, intermedium, ulnare, ?pisiform, semilunate carpal, distal carpal III, manus including metacarpal I, phalanx II-1 (13.5 mm), phalanx II-2 (18.53 mm), metacarpal III (22.49 mm), phalanx III-1 (10.58 mm), phalanx III-2 (10.54 mm), phalanx III-3 (11.5 mm), 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)
(AMNH 27435) (two or three individuals) material including carpals, distal carpal IV, metacarpal I (13.3 mm), phalanx I-1 (20.5 mm), manual ungual I (21.1 mm), metacarpal II (24.7 mm), phalanx II-1 (16 mm), phalanx II-2 (21.1 mm), manual ungual II (20.7 mm), metacarpal III (27 mm), phalanx III-1 (12.8 mm), phalanx III-2 (13 mm), phalanx III-3 (15.8 mm), manual ungual III (15.9 mm), metacarpal IV (18.5 mm), phalanx IV-1 (4.8 mm) and tibia (Legendre et al., 2013)
(AMNH 30631) distal ?ulna, radiale, intermedium fused to ulnare, centrale, semilunate carpal, distal carpal III fused to distal carpal IV, metacarpal I (9.22 mm), phalanx I-1 (12.07 mm), manual ungual I (12.88 mm), metacarpal II (17.92 mm), phalanx II-1 (10.26 mm), phalanx II-2 (11.63 mm), manual ungual II (12.06 mm), metacarpal III (18.19 mm), phalanx III-1 (7.43 mm), phalanx III-2 (6.74 mm), phalanx III-3 (8.99 mm), manual ungual III (8.87 mm), metacarpal IV (9.83 mm), phalanx IV-1 (1.81 mm), metacarpal V (1.06 mm) (Xu et al., 2009)
(CM 31374) skull, mandible (Downs, 2000)
(CM C-1-82) (one or two individuals) partial skeleton (Colbert, 1989)
....(CM 81766) partial skeleton including metacarpal I (10.8 mm), phalanx I-1 (15.7 mm), manual ungual I (12.9 mm), metacarpal II (20.7 mm), phalanx II-1 (10.7 mm), phalanx II-2 (16.2 mm), manual ungual II (14.8 mm), metacarpal III (20.6 mm), phalanx III-1 (8.5 mm), phalanx III-2 (9.4 mm), phalanx III-3 (10.5 mm), metacarpal IV (16.2 mm) (Rinehart, Lucas, Heckert, Spielmann and Celeskey, 2009)
(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)
(CMNH 11892) metatarsus (Tykoski, 2005)
(CMNH 11893) metatarsus (Tykoski, 2005)
(CMNH 11894) tibia, fibula, astragalus, calcaneum (Tykoski, 2005)
(CMNH 11895) scapulocoracoid (Tykoski, 2005)
(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 radiale, ulnare, ?pisiform, semilunate carpal and metacarpal V (Colbert, 1989)
(MCZ 4330) pelvis (Colbert, 1989)
(MCZ 4331) two distal caudal vertebrae, femur (118.2 mm), tibia (149.5 mm), metatarsal III (90.1 mm), other elements (Colbert, 1989)
(MCZ 4331a) (first individual) last thirteen dorsal vertebrae, dorsal ribs, humerus (85.0 mm), radius, ulna, radiale, intermedium, ulnare, semilunate carpal, distal carpal IV, metacarpal I (8.41 mm), metacarpal II (16.1 mm), metacarpal III (18.09 mm), phalanx III-1 (6.33 mm), metacarpal IV (10.61 mm), pelvis, hindlimbs including femur (162.9 mm) and tibia (173.0 mm) (Colbert, 1989)
(MCZ 4331b) three cervical vertebrae, three dorsal vertebrae, pelvis, femora (128.7 mm), 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, two manus, pelvis, hindlimbs (Colbert, 1989)
(MCZ 4335) vertebrae (Colbert, 1989)
(MNA.V.3139) fragmentary skull, fragmentary mandibles, postcrania (Colbert, 1989)
(MNA.V.3315) skull (143 mm), mandibles, atlas, axis, third cervical vertebra (Colbert, 1989)
(MNA.V.3318) (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.V.3319) pes including metatarsal II (67 mm), metatarsal III (76 mm), metatarsal IV (69 mm) (Colbert, 1989)
(MNA.V.3320) pes including astragalus, calcaneum, metatarsal II (115 mm), metatarsal III (128 mm), metatarsal IV (118 mm) (Colbert, 1989)
(MNA.V.3321) distal tibia, astragalocalcaneum (Colbert, 1989)
(MNA.V.3322) partial skull (Colbert, 1989)
(MNA.V.3323) caudal series (Colbert, 1989)
(NMMNH P-42200) (gracile) specimen including skull (123 mm) and sclerotic ring (Rinehart, Lucas, Heckert and Hunt, 2004)
(NMMNH P-42352) skeleton including cololite and coprolite (Rinehart, Hunt, Lucas, Heckert and Smith, 2005)
(NMMNH P-42353) skeleton including furcula (Rinehart, Lucas and Hunt, 2007)
(NMMNH P-42576) skeleton including furcula and forelimbs including metacarpal V (Rinehart, Lucas and Hunt, 2007)
(NMMNH P-42577) skeleton including furcula (Rinehart, Lucas and Hunt, 2007)
(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-46615) skeleton including furcula (Rinehart, Lucas and Hunt, 2007)
(NMMNH P-C-8-82) (several individuals including three juveniles) material including five furculae (Rinehart, Lucas and Hunt, 2006)
(SMP VP-1072) femur (Sullivan and Lucas, 1999)
(TMM 45559) (at least two individuals) material including cervical vertebrae, sacral vertebrae, partial ilium, femora, tibia, astragalocalcaneum (Tykoski, 2005)
(TMP 1984.063.0023) skull (Colbert, 1989)
(TMP 1984.063.0029) skull, mandibles, cervical vertebrae, dorsal vertebrae (Colbert, 1989)
(TMP 1984.063.0030) cervical vertebrae, dorsal vertebrae, scapula, coracoid, humerus, radius, ulna, manus (Colbert, 1989)
(TMP 1984.063.0031) maxilla, mandible (Colbert, 1989)
(TMP 1984.063.0032) anterior skull, anterior mandible, posterior cervical vertebrae, dorsal vertebrae, ribs, scapulocoracoid, forelimbs (Colbert, 1989)
(TMP 1984.063.0033) 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)
(TMP 1984.063.0034) last seven dorsal vertebrae, several caudal vertebrae, pelvis, femora, tibiae, fibula, astragalocalcaneum, pes (Colbert, 1989)
(TMP 1984.063.0035) eleven caudal vertebrae, humerus(?), tibia, fibula, astragalus, calcaneum, pes (Colbert, 1989)
(TMP 1984.063.0036) vertebrae, femora, tibiae, other elements (Colbert, 1989)
(TMP 1984.063.0037) vertebrae, scapulacoracoid(?), ischia, hindlimbs (Colbert, 1989)
(TMP 1984.063.0038) pelvis (Colbert, 1989)
(TMP 1984.063.0039) vertebrae, ilium, femur (Colbert, 1989)
(TMP 1984.063.0040) manus (Colbert, 1989)
(TMP 1984.063.0041) six caudal vertebrae (Colbert, 1989)
(TMP 1984.063.0042) partial femora, tibiae (Colbert, 1989)
(TMP 1984.063.0043) seven series of caudal vertebrae (Colbert, 1989)
(TMP 1984.063.0044) vertebrae, other elements (Colbert, 1989)
(TMP 1984.063.0045) fragmentary skull (Colbert, 1989)
(TMP 1984.063.0046) manus (Colbert, 1989)
(TMP 1984.063.0047) distal tibia, distal fibula, astragalocalcaneum (Colbert, 1989)
(TMP 1984.063.0048) sacrum, posterior ilia (Colbert, 1989)
(TMP 1984.063.0049) eleven vertebrae (Colbert, 1989)
(TMP 1984.063.0050) manus (Colbert, 1989)
(TMP 1984.063.0051) dorsal vertebrae, dorsal ribs (Colbert, 1989)
(TMP 1984.063.0052) carpus, manus (Colbert, 1989)
(TMP 1984.063.0053) seven cervical vertebrae (Colbert, 1989)
(TMP 1984.063.0054) ten caudal vertebrae (Colbert, 1989)
(TMP 1984.063.0055) femur (Colbert, 1989)
(TMP 1984.063.0056) fourth distal tarsal, metatarsal (Colbert, 1989)
(TMP 1984.063.0057) vertebrae (Colbert, 1989)
(TMP 1984.063.0058) caudal vertebrae, long bone (Colbert, 1989)
(TMP 1984.063.0059) six posterior cervical vertebrae (Colbert, 1989)
(TMP 1984.063.0060) element (Colbert, 1989)
(TMP 1984.063.0061) 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)
Diagnosis- (after Tykoski, 1998) anterior pedunclar foramina in cervical vertebrae (unknown in Coelophysis rhodesiensis).
(after Downs, 2000) unspecified differences from Coelophysis rhodesiensis in cervical length, proximal and distal hindlimb proportions and proximal caudal vertebral anatomy.
(after Tykoski and Rowe, 2004) differs from Coelophysis rhodesiensis and "Syntarsus" kayentakatae in lacking pit at the base of the nasal process of the premaxilla.
(after Bristowe and Raath, 2004) differs from Coelophysis rhodesiensis 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 Coelophysis rhodesiensis in having a square-shaped rostral process of the jugal; unreduced medial condyle of the distal trochlea of the metacarpal I.
(after Barta, Nesbitt and Norell, 2018) differs from Coelophysis rhodesiensis in lacking a pisiform; metacarpal V present.
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 neotheropods- 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. [entry in progress]
Original Coelophysis- The original type material was discovered in 1881 at Arroyo Seco and described by Cope (1887a) as Coelurus bauri and C. longicollis, though they were referred to that genus without justification. 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). In 1889, Cope separated the three species from Tanystropheus as his new genus Coelophysis because the vertebrae have neural canals, although that is also true in the former genus. Huene (1906, 1915) illustrated the material and described it in more depth, but because of the numerous complete skeletons discovered at the Coelophysis Quarry in 1947, the Arroyo Seco fragments were largely ignored until the 1980s.
Ghost Ranch- On June 22 1947 an extensive bonebed of coelophysids was discovered at Ghost Ranch (later specified as the Whitaker Quarry or Coelophysis Quarry in 1989 to distinguish it from the Canjilon Quarry also near Ghost Ranch, with Coelophysis Quarry being the term generally used today) 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. Padian (1986) first articulated the issue that the Arroyo Seco elements were less diagnostic than associated skeletons like UCMP 129618 found in 1982 or the numerous Coelophysis Quarry skeletons. Hunt and Lucas (1991) attempted to solve this by naming the Coelophysis Quarry specimens Rioarribasaurus colberti, but the ICZN (1996) ruled that a Coelophysis Quarry specimen (AMNH 7224) is the neotype of Coelophysis bauri, leaving the Arroyo Seco specimens as not definitely Coelophysis. These Arroyo Seco specimens, the Petrified Forest theropod (UCMP 129618 plus a few referred specimens), the Snyder Quarry coelophysid (assigned to C. bauri by Spielmann et al., 2007), and other Late Triassic American material may be referrable to C. bauri or at least Coelophysis, but this cannot be determined until the Ghost Ranch specimens are redescribed (although the consensus is Lepidus, Camposaurus- possibly including the Hayden Quarry material, and Gojirasaurus are all distinct). The illustrations of Coelophysis bauri in Colbert (1989) are inaccurate and have hampered comparison to other coelophysids. Downs (2000) notes most of the supposed differences from C. rhodesiensis are not real (both have a vaulted palate; interdental plates; obturator fenestra; pubic fenestra; fused pelvis; triangular dorsal transverse processes; identical hindlimb morphology; the supposed nasal fenestra of rhodesiensis is just the standard saurischian posterolateral nasal process also known in bauri). 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. Wang et al. (2017) "noted a high degree of polymorphism in skeletons from that quarry traditionally referred to Coelophysis bauri, even considering ontogenetic variation" and suggested "there may be additional basal theropod taxa represented by material from Ghost Ranch (possibly Daemonosaurus)." Barta et al. (2018) noted that for the carpus "the main variable characters within the Ghost Ranch specimens of C. bauri are the separation or fusion of the intermedium and ulnare (or alternatively the presence or absence of an intermedium altogether), presence or absence of an ossified centrale, and the degree of fusion among the distal carpals."
The manus of AMNH 30631 was first photographed by Xu et al. (2009) without a specimen number as figure S3b. It was later described and illustrated in detail by Barta et al. (2018), who stated it "is from a smaller block (#36) from the 1947 excavation of the Coelophysis Quarry" and "was found with much of a foot, but this manus and pes cannot be connected to any other individual in the quarry."
Note YPM 5705, listed by Galton (1971) in Table 2 as Coelophysis, is a cast of the neotype AMNH 7224 (YPM online).
Supposed Eucoelophysis TMP 1984.063.0033- Rinehart et al. (2009) refer a partial skeleton from the Coelophysis Quarry (TMP 1984.063.0033) to Eucoelophysis. The authors list several characters supposedly showing this isn't Coelophysis bauri unlike the other thirty-three dinosaurs in the block. Four sacrals are reported, with the last not fused to the others. "The distal scapula is less rounded and the middle shaft portion is distinctly wider than in Coelophysis bauri", but Eucoelophysis has a scapular blade which is narrower than Coelophysis. The transverse proximal femoral groove is also present in coelophysids (e.g. UCMP 129618), while the anterior trochanter was said to be very similar to gracile rhodesiensis so would be expected in a gracile Coelophysis bauri. Rinehart et al. state "A small, sharp, distinct, crest-like anterolateral trochanter is located immediately anterior to the greater trochanter on the anterior surface of the femur head as is seen in Eucoelophysis baldwini (Fig. 42E-F). This trochanter is unique, and we consider it to be the single most diagnostic feature in our assignment of TMP84-63-33 to Eucoelophysis." Yet this is merely the dorsolateral trochanter, common in dinosauriforms including Coelophysis bauri itself (Nesbitt, 2011). The authors also say "The tibia shows an appressed tibia surface (a wide, shallow sulcus to accommodate the fibular shaft) as in Eucoelophysis baldwini", which would count against a neotheropod identity if true. Finally, TMP 1984.063.0033 is said to have distal tarsals "that are very much more robust than any observed in Coelophysis bauri", but distal tarsal IV is described as 22% as thick as transversely wide while that of UCMP 129618 is 42%. Thus the only potentially valid suggested differences from Coelophysis are one less fused sacral, the broader and more angled scapula, and the absent fibular crest on the tibia, none of which are clearly figured. If these are true, referral to the contemporaneous Daemonosaurus is possible if that taxon is a neotheropod based on the cervical pleurocoels, confluent supracetabular crest and brevis ridge, fused astragalocalcaneum, "attenuated first digit [of the pes], and the fifth metatarsal ... reduced to a thin splint of bone." However these characters (except for the pleurocoels) along with the elongate dorsal centra, long postacetabular process, deep brevis fossa and small pubic boot are incompatable with a herrerasaur, which Daemonosaurus has recently been argued to be. It seems most likely given the similarity to coelophysoid-grade taxa that this is merely a misinterpreted Coelophysis bauri specimen.
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.
Not Coelophysis- 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 centra from "Lot's Wife" (actually Agate Bridge N) of the Sonsela Member of the Chinle Formation of Arizona referred to Coelophysis by Colbert (1989) are Archosauromorpha indet. (Parker and Irmis, 2005). Sullivan (1994) reported "indeterminate ceratosaur fossils which are considered topotypic material of Coelophysis bauri" found in 1993 from five sites around Arroyo Seco. Sullivan et al. (1996) published the specimens and localities, again proposing them as topotypes for Coelophysis bauri. These are undescribed with only SMP VP-487 figured, and are probably indeterminate at levels between Coelophysidae and Archosauria.
References- Cope, 1887a. The dinosaurian genus Coelurus. American Naturalist. 21, 367-369.
Cope, 1887b. A contribution to the history of the Vertebrata of the Trias of North America. Proceedings of the American Philosophical Society. 24, 209-228.
Cope, 1889. On a new genus of Triassic Dinosauria. American Naturalist. 23, 626.
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.
Hay, 1930. Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution of Washington. 390(II), 1074 pp.
Colbert, 1947. The little dinosaurs of Ghost Ranch. Natural History. 56, 392-399.
Colbert and Baird, 1958. Coelurosaur bone casts from the Connecticut Valley Triassic. American Museum Novitates. 1901, 11 pp.
Colbert, 1964. The Triassic dinosaur genera Podokesaurus and Coelophysis. American Museum Novitates. 2168, 12 pp.
Galton, 1971. The prosauropod dinosaur Ammosaurus, the crocodile Protosuchus, and their bearing on the age of the Navajo Sandstone of northeastern Arizona. Journal of Paleontology. 45(5), 781-795.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
Padian, 1986. On the type material of Coelophysis Cope (Saurischia: Theropoda) and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation). In Padian (ed.). The Beginning of the Age of Dinosaurs: Faunal Change Across the Triassic-Jurassic Boundary. Cambridge University Press. 45-60.
Colbert, 1989. The Triassic dinosaur Coelophysis. Museum of Northern Arizona Bulletin. 57, 1-174.
Colbert, 1990. Variation in Coelophysis bauri. In Carpenter and Currie (eds.). Dinosaur Systematics. Approaches and Perspectives. Cambridge University Press. 81-90.
Rowe and Gauthier, 1990. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 151-168.
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Colbert, Harris, Charig, Dodson, Gillette, Ostrom and Weishampel, 1992. Coelurus bauri Cope, 1887 (currently Coelophysis bauri; Reptilia, Saurischia): Proposed replacement of the lectotype by a neotype. Bulletin of Zoological Nomenclature. 49, 276-279.
Creisler, 1993. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(4), 292-294.
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Holtz, 1993. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(3), 237-238.
Hotton, 1993. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(4), 294.
Hunt and Lucas, 1993. Comments on a proposed neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(2), 147-150.
Jenkins, 1993. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(3), 238-239.
Molnar, 1993. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(3), 236-237.
Paul, 1993. Are Syntarsus and the Whitaker quarry theropod the same genus? New Mexico Museum of Natural History and Science Bulletin. 3, 397-402.
Schwartz, 1993. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(3), 236.
Spinar, 1993. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(3), 237.
Sullivan, 1993. Comments on a proposed neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(2), 150-151.
Tubbs, 1993. Comments on a proposed neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 50(2),
Glut, 1994. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(1), 50.
Huber, 1994. Comment on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(2), 156-158.
Jacobs, 1994. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(1), 50.
Lucas and Hunt, 1994. Comment on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(3), 265-266.
Nicholls, 1994. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(1), 50.
Olshevsky, 1994. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(1), 49-50.
de Ricqles, 1994. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(1), 51.
Sullivan, 1994. Topotypic material of Coelophysis bauri (Cope) and the Coelophysis-Rioarribasaurus-Syntarsus problem. Journal of Vertebrate of Paleontology. 14(3), 48A.
Tubbs, 1994. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(1), 51.
Welles, 1994. Comments on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 51(1), 48.
Sullivan, 1995. Comment on the proposed designation of a neotype for Coelophysis bauri (Cope, 1887) (Reptilia, Saurischia). Bulletin of Zoological Nomenclature. 52, 76-77.
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.
Smith, 1997. Cranial allometry in Coelophysis: Preliminary results. Southwest Paleontological Symposium- Proceedings. 41-49.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the western United States. Geobios. 31(4), 511-531.
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.
Downs, 2000. Coelophysis bauri and Syntarsus rhodesiensis compared, with comments on the perparation and preservation of fossils from the Ghost Ranch Coelophysis Quarry. New Mexico Museum of Natural History and Science Bulletin. 17, 33-37.
Smith, Andersen, Larsson and Bybee, 2000. Results of a high resolution CT-scan of Coelophysis bauri (Upper Triassic Chinle Formation, New Mexico). Journal of Vertebrate Paleontology. 20(3), 70A.
Rinehart, Lucas and Heckert, 2001. Preliminary statistical analysis defining the juvenile, robust and gracile forms of the Triassic dinosaur Coelophysis. Journal of Vertebrate Paleontology. 21(3), 93A.
Gay, 2002. The myth of cannibalism in Coelophysis bauri. Journal of Vertebrate Paleontology. 22(3), 57A.
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.
Rinehart, Lucas, Heckert and Hunt, 2004. Vision characteristics of Coelophysis bauri based on sclerotic ring, orbit, and skull morphology. Journal of Vertebrate Paleontology. 24(3), 207A.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 47-70.
Lucas, Sullivan, Hunt and Heckert, 2005. The saga of Coelophysis. 56th NMGS Fall Field Conference 2004, First-day Road Log. 37-38.
Parker and Irmis, 2005. Advances in Late Triassic vertebrate paleontology based on new material from Petrified Forest National Park, Arizona. New Mexico Museum of Natural History and Science Bulletin. 29, 45-58.
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 thesis. 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, 2006a. Digesting the Coelophysis-cannibal hypothesis and its importance to prey choice in theropod dinosaurs. Journal of Vertebrate Paleontology. 26(3), 105A.
Nesbitt, Turner, Erickson and Norell, 2006b. 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 (online 2006). The cranial anatomy of the coelophysoid theropod Zupaysaurus rougieri from the Upper Triassic of Argentina. Historical Biology. 19(2), 185-202.
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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, Heckert, Lucas and Celeskey, 2008. Growth, allometry, and age/size distribution of the Late Triassic theropod dinosaur Coelophysis bauri: Preliminary results. Journal of Vertebrate Paleontology. 28(3), 132A.
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C? rhodesiensis (Raath, 1969) Paul, 1988
= Syntarsus rhodesiensis Raath, 1969
= Megapnosaurus rhodesiensis (Raath, 1969) Ivie, Slipinski and Wegrzynowicz, 2001
Hettangian-Sinemurian, Early Jurassic
Southcote Farm, Forest Sandstone Formation, Zimbabwe
Holotype- (QG 1) (2.02 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 (129 mm), humerus (100 mm), radius (61 mm), ulna (77 mm), radiale, intermedium, ulnare, pisiform, semilunate carpal, distal carpal III, metacarpal I (12.66 mm), phalanx I-1 (20 mm), manual ungual I (18 mm), metacarpal II (25.37 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 (17.4 mm), phalanx IV-1 (4.13 mm), ilia (one partial; 148 mm), pubis (204 mm), ischia (130 mm), femur (208 mm), tibiae (223 mm), fibulae (one partial; 208 mm), astragalocalcanea (29 mm wide), distal tarsals IV, metatarsals I (30 mm), phalanx I-1 (17 mm), pedal ungual I (14 mm), metatarsals II (119 mm), phalanges II-1 (33 mm), phalanges II-2 (one proximal; 25 mm), pedal ungual II (22 mm), metatarsals III (132 mm), phalanges III-1 (37 mm), phalanges III-2 (29 mm), phalanges III-3 (24 mm), pedal ungual III (22 mm), metatarsals IV (117 mm), phalanges 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- (QG 3A) (robust) lacrimal, squamosal, dentaries, cervical vertebrae, dorsal vertebrae, ribs, sacral vertebrae, caudal vertebrae, chevrons, proximal femora, proximal tibia (Raath, 1977)
Hettangian-Sinemurian, Early Jurassic
Maujra River, Forest Sandstone Formation, Zimbabwe
(QG 45) (gracile juvenile) sacrum, ilia, femora, tibia, pes, fragments (Raath, 1977)
(QG 76) (gracile adult) femur (201 mm) (Raath, 1977)
Hettangian-Sinemurian, Early Jurassic
Chitake River, Forest Sandstone Formation, Zimbabwe
(QG numbers below) including those elements listed below, at least 26 individuals are present, represented by 21 premaxillae, 31 maxillae, 13 nasals, 13 lacrimals, 7 prefrontals, 19 frontals, 21 parietals, 10 postorbitals, 11 squamosals, 2 jugals, 3 quadratojugals, 19 quadrates, 5 braincases, 6 palatines, 8 pterygoids, 6 ectopterygoids, a few sclerotic plates, 21 dentaries, 11 splenials, coronoid, 11 surangulars, 15 angulars, 11 prearticulars, 18 articulars, many teeth, 77 cervical vertebrae, many more than 7 cervical ribs, 36 dorsal vertebrae, more than 38 dorsal ribs, 7 sacral centra, 120 caudal vertebrae, 45 caudal centra, many more than 37 chevrons, 22 scapulocoracoids, 22 humeri, 10 radii, 8 ulnae, 8 manus, many manual elements, more than 10 pelves with sacra, more than 34 femora, more than 23 tibiae, more than 18 fibulae, more than 12 astragalocalcanea, 23 distal tarsals IV, many pedal elements, 12 blocks of unprepared material (Raath, 1977)
(QG 124) posterior mandible (Raath, 1977)
(QG 164) (juvenile) metatarsal I, pedal digit I, incomplete metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, incomplete metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, incomplete metatarsal IV, phalanx IV-3, phalanx IV-4, pedal ungual IV (Raath, 1977)
(QG 165) maxilla, nasals, lacrimal, jugal, quadratojugal, palatine, ectopterygoid, pterygoid, hyoids, postcrania (Raath, 1977)
(QG 169) seventh cervical vertebra (30 mm), eighth cervical vertebra (25 mm), ninth cervical vertebra (20.5 mm), tenth cervical vertebra (18 mm), first dorsal vertebra (20 mm), second dorsal vertebra (20 mm), third dorsal vertebra (23 mm), fourth dorsal vertebra (23.5 mm), fifth dorsal vertebra (25.5 mm) (Raath, 1977)
(QG 170) ~eighth cervical vertebra (35 mm), ~ninth cervical vertebra (28 mm) (Raath, 1977)
(QG 171) incomplete second dorsal vertebra (26.5 mm), incomplete third dorsal vertebra (27 mm) (Raath, 1977)
(QG 172) partial fifth cervical vertebra, sixth cervical vertebra (37 mm), seventh cervical vertebra (33 mm), eighth cervical vertebra (27 mm), ninth cervical vertebra (22 mm) (Raath, 1977)
(QG 173) sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth cervical vertebra, partial cervical ribs, first dorsal vertebra , second dorsal vertebra (Raath, 1977)
(QG 174) axial neural arch, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, partial sixth cervical vertebra (Raath, 1977)
(QG 175) third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra (Raath, 1977)
(QG 176) atlas, axis (27 mm), third cervical vertebra (36 mm), fourth cervical vertebra (40 mm), partial cervical ribs (Raath, 1977)
(QG 177) atlas, axis (20 mm), third cervical vertebra (26 mm), fourth cervical vertebra (32 mm), fifth cervical vertebra (37 mm) (Raath, 1977)
(QG 178) atlas, axial neural arch, third cervical vertebra (25.5 mm) (Raath, 1977)
(QG 179) atlantal neural arch, partial axis, third cervical vertebra (33 mm), fourth cervical vertebra (36.5 mm), fifth cervical vertebra (41 mm), sixth cervical vertebra (44 mm)(Raath, 1977)
(QG 180) second dorsal vertebra (22 mm) (Raath, 1977)
(QG 181-192) (Raath, 1977)
(QG 193) maxilla, nasal, lacrimal, jugal, prefrontal, frontals, parietals, postorbital, squamosal, quadratojugal, braincase, mandible, scapulocoracoid (157 mm), two furculae (Raath, 1977)
(QG 194) maxilla, nasals, squamosal, quadratojugal, quadrate, braincase, ectopterygoid, pterygoid (Raath, 1977)
(QG 195) braincase (Raath, 1977)
(QG 196) braincase (Raath, 1977)
(QG 197) braincase (Raath, 1977)
(QG 198-201) (Raath, 1977)
(QG 202) premaxillae, incomplete maxilla, partial nasal, incomplete dentaries (Raath, 1977)
(QG 203-205) (Raath, 1977)
(QG 206) maxilla (Raath, 1977)
(QG 207) (Raath, 1977)
(QG 208) maxilla, three sclerotic plates, partial mandible (Raath, 1977)
(QG 209) maxilla (Raath, 1977)
(QG 210) maxilla (Raath, 1977)
(QG 211) (Raath, 1977)
(QG 212) maxilla (Raath, 1977)
(QG 213) maxilla (Raath, 1977)
(QG 214-229) (Raath, 1977)
(QG 230) partial lacrimal (Raath, 1977)
(QG 231-233) (Raath, 1977)
(QG 234) lacrimal (Raath, 1977)
(QG 235) quadratojugal, quadrate, palatine, ectopterygoid, pterygoid (Raath, 1977)
(QG 236-240) (Raath, 1977)
(QG 241) palatine, ectopterygoid, pterygoid (Raath, 1977)
(QG 242-243) (Raath, 1977)
(QG 244) furcula (Raath, 1977)
(QG 245) premaxilla, premaxillary teeth (Raath, 1977)
(QG 246) premaxillae, premaxillary teeth (Raath, 1977)
(QG 247-248) (Raath, 1977)
(QG 249) premaxilla (Raath, 1977)
(QG 250-253) (Raath, 1977)
(QG 254) premaxilla (Raath, 1977)
(QG 255-262) (Raath, 1977)
(QG 263) pterygoid (Raath, 1977)
(QG 264) (Raath, 1977)
(QG 265) pterygoid (Raath, 1977)
(QG 266-277) (Raath, 1977)
(QG 278) incomplete jugal, lacrimal, prefrontal (Raath, 1977)
(QG 279-286) (Raath, 1977)
(QG 287) postorbital (Raath, 1977)
(QG 288-302) (Raath, 1977)
(QG 303) dentary (Raath, 1977)
(QG 304) (Raath, 1977)
(QG 305) dentary (Raath, 1977)
(QG 306) (Raath, 1977)
(QG 307) mandible (Raath, 1977)
(QG 308-395) (Raath, 1977)
(QG 396) first dorsal centrum (23.5 mm) (Raath, 1977)
(QG 397-404) (Raath, 1977)
(QG 405) sixth dorsal vertebra (29 mm) (Raath, 1977)
(QG 406) fifth dorsal vertebra (29 mm) (Raath, 1977)
(QG 407) (Raath, 1977)
(QG 408) partial sixth dorsal vertebra, seventh dorsal vertebra (27.5 mm), eighth dorsal neural arch (Raath, 1977)
(QG 409-412) (Raath, 1977)
(QG 413) incomplete posterior dorsal vertebra (26.5 mm), incomplete posterior dorsal vertebra (25 mm) (Raath, 1977)
(QG 414-422) (Raath, 1977)
(QG 423) partial cervical ribs (Raath, 1977)
(QG 424-508) (Raath, 1977)
(QG 509) mid chevron (Raath, 1977)
(QG 510) distal chevron (Raath, 1977)
(QG 511) (Raath, 1977)
(QG 512) scapulocoracoid (121 mm) (Raath, 1977)
(QG 513) (Raath, 1977)
(QG 514) (robust) scapulocoracoid, humerus (102 mm), radius (62.5 mm), ulna (71 mm) (Raath, 1977)
(QG 514b) (robust) ulna (Raath, 1977)
(QG 515-516) (Raath, 1977)
(QG 517) scapula (80 mm), humerus (74.5 mm) (Raath, 1977)
(QG 518-523) (Raath, 1977)
(QG 524) partial scapula, humerus (87 mm) (Raath, 1977)
(QG 525-544) (Raath, 1977)
(QG 545) (gracile) humerus (Raath, 1977)
(QG 546-549) (Raath, 1977)
(QG 550) humerus (76.5 mm) (Raath, 1977)
(QG 551-562) (Raath, 1977)
(QG 563) radius (54 mm), ulna (60 mm) (Raath, 1977)
(QG 564-567) (Raath, 1977)
(QG 568) (gracile) ulna (Raath, 1977)
(QG 569-572) (Raath, 1977)
(QG 573) (robust) metacarpals I (14.08, 14.69 mm), phalanx I-1 (18.99 mm), manual ungual I (22.61 mm), metacarpals II (28.25, 29.96 mm), phalanx II-1 (16.93 mm), phalanges II-2 (21.48, 21.54 mm), manual ungual II or III (18 mm), metacarpals III (31.67, 30.5 mm), phalanx III-1 (14.09 mm), phalanx III-2 (11.22 mm), metacarpals IV (21.77 mm) (Raath, 1977)
(QG 574-576) (Raath, 1977)
(QG 577) (gracile) distal radius, distal ulna, metacarpal I (12.09 mm), phalanx I-1 (16.98 mm), manual ungual I (14.4 mm), metacarpal II (24.18 mm), phalanx II-1 (12.25 mm), phalanx II-2 (17.44 mm), manual ungual II (6.09 mm), metacarpal III (24.44 mm), phalanx III-1 (9.14 mm), phalanx III-2 (9.41 mm), phalanx III-3 (12.04 mm), manual ungual III (6.99 mm), metacarpal IV (14.35 mm), phalanx IV-1 (3.95 mm) (Raath, 1977)
(QG 578-685) (Raath, 1977)
(QG 686) partial forelimb including metacarpal I (12.54 mm), metacarpal II (26.78 mm), proximal phalanx II-1, metacarpal III (27.68 mm), proximal metacarpal IV (Raath, 1977)
(QG 687) distal carpal (Raath, 1977)
(QG 688-690) (Raath, 1977)
(QG 691) (gracile juvenile) ilium (122 mm), incomplete pubis, ischium (125.5 mm), femora (142 mm), tibia (156 mm), fibula (152 mm), astragalus, calcaneum (Raath, 1977)
(QG 692-695) (Raath, 1977)
(QG 696) ilium (148 mm), proximal pubis, (Raath, 1977)
(QG 697-712) (Raath, 1977)
(QG 713) (gracile juvenile) proximal femur (Raath, 1977)
(QG 714) (subadult) femoral fragment (Raath, 1977)
(QG 715) (gracile juvenile) incomplete femur (Raath, 1977)
(QG 716) (rubust subadult) proximal femur (Raath, 1977)
(QG 717) (gracile adult) proximal femur (Raath, 1977)
(QG 718-721) (Raath, 1977)
(QG 722) distal femur (Raath, 1977)
(QG 723) (juvenile) distal femur (Raath, 1977)
(QG 724) (Raath, 1977)
(QG 725) (robust adult) proximal femur (Raath, 1977)
(QG 726) (robust adult) incomplete femur (Raath, 1977)
(QG 727) (robust adult) incomplete femur (Raath, 1977)
(QG 728) (Raath, 1977)
(QG 729) (robust adult) proximal femur (Raath, 1977)
(QG 730) (Raath, 1977)
(QG 731) (robust subadult) femur (189 mm) (Raath, 1977)
(QG 732) (robust adult) proximal femur (Raath, 1977)
(QG 733) (robust adult) proximal femur (Raath, 1977)
(QG 734) distal femur (Raath, 1977)
(QG 735-737) (Raath, 1977)
(QG 738) (gracile subadult) femur (192 mm) (Raath, 1977)
(QG 739) (gracile adult) proximal femur (Raath, 1977)
(QG 740) (gracile adult) proximal femur (Raath, 1977)
(QG 741) (Raath, 1977)
(QG 742) (gracile adult) proximal femur (Raath, 1977)
(QG 743) (adult) femoral fragment (Raath, 1977)
(QG 744) (gracile juvenile) proximal femur (Raath, 1977)
(QG 745) (gracile juvenile) femur (172 mm) (Raath, 1977)
(QG 746) femoral fragment (Raath, 1977)
(QG 747) (juvenile) distal femur (Raath, 1977)
(QG 748) partial femur (Raath, 1977)
(QG 749-752) (Raath, 1977)
(QG 753) (robust adult) proximal femur (Raath, 1977)
(QG 754) (robust subadult) femur (186 mm) (Raath, 1977)
(QG 755) (robust adult) femur (185 mm) (Raath, 1977)
(QG 756) (subadult?) proximal femur (Raath, 1977)
(QG 757) distal femur (Raath, 1977)
(QG 758-759) (Raath, 1977)
(QG 760) (robust adult) proximal femur (Raath, 1977)
(QG 761) (Raath, 1977)
(QG 762) tibia (179 mm), proximal fibula (Raath, 1977)
(QG 763-767) (Raath, 1977)
(QG 768) (gracile juvenile) distal tibia, distal fibula, astragalus, calcaneum, distal tarsal III, distal tarsal IV, proximal metatarsal II, proximal metatarsal III, proximal metatarsal IV (Raath, 1977)
(QG 769) (Raath, 1977)
(QG 770) tibiotarsus (210 mm), fibula (197 mm) (Raath, 1977)
(QG 771-780) (Raath, 1977)
(QG 781) astragalus, calcaneum (Raath, 1977)
(QG 782-784) (Raath, 1977)
(QG 785) astragalus (Raath, 1977)
(QG 786) (gracile) astragalus, calcaneum (Raath, 1977)
(QG 787-802) (Raath, 1977)
(QG 803) tibia (205), fibula (194 mm), astragalocalcaneum (33 mm trans) (Raath, 1977)
(QG 804) (Raath, 1977)
(QG 805) tibia (207 mm), fibula (201 mm), astragalocalcaneum (31 mm trans) (Raath, 1977)
(QG 806-815) (Raath, 1977)
(QG 816) astragalus, calcaneum (Raath, 1977)
(QG 817-823) (Raath, 1977)
(QG 824) distal tarsal IV (Raath, 1977)
(QG 825) distal tarsal IV (Raath, 1977)
(QG 826-829) (Raath, 1977)
(QG 830) distal tarsal IV (Raath, 1977)
(QG 831) two distal tarsals IV (Raath, 1977)
(QG 832-845) (Raath, 1977)
(QG 846) distal tarsal III (Raath, 1977)
(QG 847) (Raath, 1977)
(QG 848) distal tarsal III (Raath, 1977)
(QG 849-850) (Raath, 1977)
(QG 851) distal tarsal III (Raath, 1977)
(QG 852) (Raath, 1977)
(QG 853) distal tarsal III) (Raath, 1977)
(QG 854-1103) (Raath, 1977)
hundreds of elements including skull, cranial material, cervical vertebrae and parts of all portions of the skeleton (Roberts et al., 2008)
Pliensbachian, Early Jurassic
Edelweiss/Welbedacht Farms, Upper Elliot Formation, South Africa
(BPI/1/5246) partial ilium (Munyikwa and Raath, 1999)
Pliensbachian, Early Jurassic
Mequatling Farm, Upper Elliot Formation, South Africa
(BPI coll.; = field numbers F7, F43) (at least eight individuals) cervical vertebra, dorsal vertebra, several distal femora, proximal tibia, pedal fragments (Raath 1980)
Pliensbachian, Early Jurassic
Spioenkop Farm, Upper Elliot Formation, South Africa
(BPI/1/coll.) partial skeleton (Blackbeard and Yates, 2007)
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 "Syntarsus" 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- The holotype was discovered in 1963, and the large bonebed from the Chitake River (QG 124, 164-165, 169-1103) was found in 1972. 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. 4(28), 1-25.
Galton, 1971. Manus movements of the coelurosaurian dinosaur Syntarsus and opposability of the theropod hallux. Arnoldia. 5(15), 1-8.
Raath, 1977. The anatomy of the Triassic theropod Syntarsus rhodesiensis (Saurischia: Podokesauridae) and a consideration of its biology. PhD thesis. Rhodes University. 233 pp.
Raath, 1980. The theropod dinosaur Syntarsus (Saurischia: Podokesauridae) discovered in South Africa. South African Journal of Science. 76(8), 375-376.
Raath, 1985. The theropod Syntarsus and its bearing on the origin of birds. In Hecht, Ostrom, Viohl and Wellnhofer (eds.). The Beginnings of Birds. Freunde des Jura-Museums Eichstätt, Eichstätt. 219-227.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Raath, 1990. Morphological variation in small theropods and its meaning in systematics: Evidence from Syntarsus rhodesiensis. In Carpenter and Currie (eds.). Dinosaur Systematics: Approaches and Perspectives. Cambridge University Press, Cambridge. 91-105.
Chinsamy, 1990. Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda). Palaeontologica Africana. 27, 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.
Starck and Chinsamy, 2002. Bone microstructure and developmental plasticity in birds and other dinosaurs. Journal of Morphology. 254, 232-246.
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 (eds.). The Dinosauria Second Edition. University of California Press. 47-70.
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.
Blackbeard and Yates, 2007. The taphonomy of an Early Jurassic dinosaur bonebed in the Northern Free State (South Africa). Journal of Vertebrate Paleontology. 27(3), 49A.
Ezcurra, 2007 (online 2006). The cranial anatomy of the coelophysoid theropod Zupaysaurus rougieri from the Upper Triassic of Argentina. Historical Biology. 19(2), 185-202.
Roberts, Mgodi, Broderick, Yates and O'Connor, 2008. Paleontology and taphonomy of a spectacular Late Triassic-Early Jurassic theropod bone bed ("Syntarsus" rhodesiensis) from the Zambezi Valley, Zimbabwe. Journal of Vertebrate Paleontology. 29(3), 133A.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Barta, Nesbitt and Norell, 2016. The manus of early theropod dinosaurs: Dholigital reconstructions of articulated specimens reveal a complex evolutionary pattern of digit reduction. Journal of Vertebrate Paleontology. Program and Abstracts, 93.
Barta, Nesbitt and Norell, 2018 (online 2017). The evolution of the manus of early theropod dinosaurs is characterized by high inter- and intraspecific variation. Journal of Anatomy. 232(1), 80-104.

Segisaurus Camp, 1936
S. halli Camp, 1936
Hettangian-Sinemurian, Early Jurassic
Keet Seel UCMP V3308, 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)) is enforced, Segisaurus is placed outside the Liliensternus + Coelophysis 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. Proceedings of the Geological Society of America. 1934, 384-385.
Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of Arizona. University of California Publications in Geological Sciences. 24(2), 39-56.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Senter and Hutchinson, 2001. New information on the skeleton of the theropod Segisaurus halli. Journal of Vertebrate Paleontology. 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.

unnamed clade (Passer domesticus <- Coelophysis bauri)
= Ceratosauroidea sensu Sereno, 1998
Definition- (Carnotaurus sastrei <- Coelophysis bauri) (modified)
= Neoceratosauria sensu Padian, Hutchinson and Holtz, 1999
Definition- (Ceratosaurus nasicornis <- Coelophysis bauri) (modified)
= Averostra sensu Dal Sasso, Maganuco and Cau, 2018
Definition- (Ceratosaurus nasicornis, Vultur gryphus <- Coelophysis bauri)
Diagnosis- (suggested) anteroposterior lateral jugal ridge absent; vagus foramen exits posterior to metotic strut; anterior dorsal vertebrae with ventral keel; narrow notch between preacetabular process and pubic peduncle; tibia equal to shorter than femur.
References- 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.
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.
Dal Sasso, Maganuco and Cau, 2018. The oldest ceratosaurian (Dinosauria: Theropoda), from the Lower Jurassic of Italy, sheds light on the evolution of the three-fingered hand of birds. PeerJ. 6:e5976.

unnamed Neotheropoda (Long and Murry, 1995)
Late Norian, Late Triassic
Piedra Lumbre, Petrified Forest Member of the Chinle Formation, New Mexico, US
?(MCZ 3038) metatarsal II (Nesbitt and Stocker, 2008)
(MCZ 3039) incomplete cervical vertebra (Nesbitt and Stocker, 2008)
?(MCZ 4779) distal femur (Nesbitt and Stocker, 2008)
Comments- Nesbitt and Stocker (2008) figure three specimens found in 1934 or 1935 and refer them to Theropoda. MCZ 3038 is a metatarsal II stated to be "identical to that of Coelophysis bauri and other early theropods, and fragments of an identical metatarsal II (part of UCMP 152645) were found at the Canjilon Quarry. However, no unique character states are shared between MCZ 3038 and Coelophysis bauri exclusive of other theropods." MCZ 3039 is a cervical with anterior and posterior pleurocoels, leading the authors to say that this "clearly indicates that the cervical can be assigned to a theropod, and more specifically, a coelophysoid", but more recent phylogenies place some taxa with double pleurocoels such as Dilophosaurus outside Coelophysoidea. Finally, the distal femur MCZ 4779 is said to have "a short, posteriorly pointed crista tibiofibularis, a cleft separating the crista tibiofibularis from the lateral condyle of the femur and a gently rounded lateral condyle of the femur. All of these character states are present in early theropods as well as other basal saurischians and are not diagnostic to a specific taxon." More specifically, the obtuse angle between the lateral condyle and crista tibiofibularis and the groove running posterolaterally into it are saurischian characters. Comparing it to non-averostran neotheropods, it is most similar to Liliensternus and Zupaysaurus based on the lack of an extensor groove and short transverse width compared to anteroposterior depth. The proximal outline of metatarsal II is also more similar Liliensternus, differing from the more semicircular shape of coelophysids or the short anterior edge with rounded anterolateral corner of Dilophosaurus.
Reference- Nesbitt and Stocker, 2008. The vertebrate assemblage of the Late Triassic Canjilon Quarry (northern New Mexico, USA), and the importance of apomorphy-based assemblage comparisons. Journal of Vertebrate Paleontology. 28(4), 1063-1072.

unnamed neotheropod (Benton, Martill and Taylor, 1995)
Sinemurian, Early Jurassic
Broadford Beds Formation, Scotland
Material- (NMS.G.1994.10.1) incomplete tibia
Comments- Collected in 1992, Benton et al. (1995) described this as a theropod based on the "fibula closely appressed to tibia and attached to a tibial crest" and "thin-walled, hollow, long bones", and probably a ceratosaur (sensu lato) based on its age and sharing a "straight tibia with a broad proximal end, priminent cnemial crest, and sharp fibular facet" with Coelophysis. As noted by Carrano and Sampson (2004), Benton et al. misidentified this distal left tibia as a proximal right tibia. Thus the fibular crest is unpreserved and the feature Benton et al. described is only a sharp lateral edge to the shaft, while their cnemial crest is the proximomedial buttress. Ironically, it is still very similar to Coelophysis so was correctly identified for mostly incorrect reasons, as Carrano and Sampson concluded it "cannot be identified more specifically than Coelophysoidea indet.." The distal outline appears more similar to Zupaysaurus and Liliensternus than Coelophysis based on the longer posterolateral process.
References- Benton, Martill and Taylor, 1995. The first Lower Jurassic dinosaur from Scotland: Limb bone of a ceratosaur theropod from Skye. Scottish Journal of Geology. 31(2), 177-182.
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.

Liliensternus Welles, 1984
L. liliensterni (Huene, 1934) Welles, 1984
= Halticosaurus liliensterni Huene, 1934
Late Norian, Late Triassic
Trossingen Formation, Thüringen, Germany
Syntypes- (MB.R.2175; = HMN R1291) (two gracile subadults) (5.15 m, 127 kg) (partial skull ~395 mm) three partial maxillae, partial jugal, incomplete postorbital, incomplete squamosal, quadrate (85 mm), three ectopterygoids, two ?pterygoid fragments, partial mandible, two incomplete dentaries, splenials, cervical rib fragments, four proximal dorsal ribs, dorsal rib fragments, gastralia, chevron fragments, two scapulae (one incomplete, one partial; ~30 mm), humerus, distal carpal I+II, proximal metacarpal I, distal metacarpal I, proximal phalanx I-1, metacarpal II (68 mm), proximal metacarpal II, phalanges II-2 (42 mm), incomplete manual ungual II (~30 mm), proximal metacarpal III, phalanx III-1 (42 mm), incomplete manual ungual III (~20 mm), proximal phalanx IV-1, three incomplete ilia (270 mm), phalanx II-1 (83 mm), phalanx II-2 (40 mm), incomplete pedal ungual II, phalanx III-1 (83 mm), phalanx III-2 (47 mm), phalanx III-3 (42 mm), pedal ungual III (38 mm), phalanx IV-1 (47 mm), phalanx IV-2 (32 mm), phalanx IV-3 (29 mm), phalanx IV-4 (22 mm), five pedal unguals
....(large individual) partial axis, incomplete third cervical vertebra (~80 mm), fourth cervical vertebra, incomplete fifth cervical vertebra (80 mm), sixth cervical vertebra (86 mm), eighth cervical vertebra (75 mm), ninth cervical vertebra (64 mm), tenth cervical vertebra (70 mm), two partial mid dorsal centra, incomplete twelfth dorsal vertebra (71 mm), incomplete thirteenth dorsal vertebra (70 mm), proximal first dorsal rib, second sacral vertebra (57 mm), third sacral vertebra (54 mm), sixth caudal vertebra, seventh caudal vertebra (53 mm), eighth caudal vertebra (56 mm), ninth caudal vertebra (58 mm), tenth caudal vertebra (58 mm), twelfth caudal vertebra (58 mm), thirteenth caudal vertebra (58 mm), fourteenth caudal vertebra (55 mm), twenty-fifth caudal vertebra (62 mm), twenty-sixth caudal vertebra (62 mm), twenty-seventh caudal vertebra (62 mm), twenty-eighth caudal vertebra (62 mm), twenty-ninth caudal vertebra (62 mm), thirtieth caudal vertebra (62 mm), thirty-first caudal vertebra (62 mm), thirty-second caudal vertebra (62 mm), thirty-third caudal vertebra (62 mm), thirty-fourth caudal vertebra (72 mm), thirty-fifth caudal vertebra (72 mm), thirty-sixth caudal vertebra (72 mm), thirty-seventh caudal vertebra (72 mm), thirty-eighth caudal vertebra (72 mm), partial coracoids, humerus (205 mm), radius (150 mm), ulna (155 mm), partial pubes, proximal ischium, femora (420 mm), tibiae (400 mm), incomplete fibulae, astragalocalcanea (60 mm trans), metatarsal II (205 mm), metatarsal III (220 mm), metatarsal IV (200 mm)
....(small individual) sixth cervical vertebra, eighth cervical vertebra, first dorsal vertebra (56 mm), second dorsal vertebra (50 mm), incomplete third dorsal vertebra, partial fourth neural arch, mid dorsal centrum (50 mm), mid dorsal centrum (50 mm), eleventh dorsal vertebra (55 mm), first sacral centrum (53 mm), fourth sacral vertebra (49 mm), fifth sacral vertebra (51 mm), first caudal vertebra (51 mm), second caudal vertebra (50 mm), third caudal vertebra (50 mm), fourth caudal vertebra (50 mm), fifth caudal vertebra (50 mm), sixth caudal vertebra (50 mm), eighth caudal vertebra (50 mm), ninth caudal vertebra (50 mm), tenth caudal vertebra (50 mm), twelfth caudal vertebra (52 mm), thirteenth caudal vertebra (53 mm), fourteenth caudal vertebra (53 mm), fiftheenth caudal vertebra (53 mm), sixteenth caudal vertebra (53 mm), seventeenth caudal vertebra (53 mm), eighteenth caudal vertebra (53 mm), nineteenth caudal vertebra (53 mm), twentieth caudal vertebra (53 mm), twenty-first caudal vertebra (53 mm), twenty-second caudal vertebra (53 mm), twenty-third caudal vertebra (53 mm), twenty-fourth caudal vertebra (53 mm), twenty-fifth caudal vertebra (53 mm), twenty-sixth caudal vertebra (53 mm), twenty-seventh caudal vertebra (53 mm), partial coracoids, humerus (190 mm), pubes (410 mm), incomplete ischia (250, 270 mm), femora (400 mm), tibiae (370 mm), incomplete fibulae, astragalocalcanea (60 mm trans), metatarsal II (185 mm), metatarsal III (205 mm), metatarsal IV (180 mm)
Late Norian, Late Triassic
Trossingen Formation, Heroldsberg, Germany
Referred- (UA coll.; lost) proximal metatarsal II (Meyer, 1855)
Late Norian, Late Triassic
Trossingen Formation, Halberstadt, Germany
?(MB Fund. Nr. IV in part; lost?) two teeth (Jaekel, 1914)
?(MB coll?) teeth (Jaekel, 1914)
?(MHH coll.) ?twentieth caudal centrum (14 mm), proximal tibia two metatarsals, two phalanges (Kuhn, 1939)
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 syntypes were discovered in 1932. Huene (1932) assumed nine cervicals and fourteen dorsals as compared to the modern neotheropod default of ten cervicals and thirteen dorsals. He also only identified three sacrals, but comparison to Dilophosaurus indicates his first caudal is a fourth sacral (based on the expanded transverse processes), and his second caudal would then be a fifth sacral. The metacarpals III and IV of Huene are II and III based on comparison to Dilophosaurus, which results in all of the manual material being from the next medial digit than Huene's identification (e.g. supposed proximal metacarpal V is metacarpal IV). 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 lectotype). 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.
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 "of a theropod dinosaur such as the ceratosaurian Liliensternus liliensterni" by Galton (2000) and a proximal metatarsal II of cf. Liliensternus sp. by Moser (2003). Comparison with figures of Liliensternus indicates Moser is correct, who also stated it is "the same size and shape, if not completely identical" to Liliensternus. He further notes the element is missing from the UA collections.
Jaekel (1914) reported Fund. Nr. IV as a small dinosaur with theropod-type teeth (ziphodont and finely serrated) and postcrania consisting of "a femur, an os pubis, an ilium, several vertebrae of the sacral and tail region." Sander (1992) mentioned it as "theropod postcranial remains which were briefly described by Jaekel (1914a, p. 195) as indeterminate carnosaur" which he "listed as pertaining to cf. Liliensternus." However, Huene (1932) described the postcrania as cf. Palaeosaurus (?) sp. (aff. diagnosticus), which by that time consisted only of a posterior dorsal centrum, two partial distal caudal vertebrae, a partial ilium and incomplete pubis. It shows the squared postacetabular process and hypertrophied semilunate pubic tubercle Yates found diagnostic of Efraasia minor, so is here referred to that taxon. It is unknown whether the theropod teeth were lost or recatalogued. Jaekel also noted "several [Plateosaurus] skeletons were accompanied by broken-off tooth crowns of carnosaurs", but these teeth remain undescribed. Sander referred Pterospondylus from these same deposits to Liliensternus as well, but as noted by Rauhut and Hungerbuhler "the transverse processes in ... Liliensternus are less strongly backturned and not as significantly triangular" as Pterospondylus.
Kuhn (1939) referred several elements to Halticosaurus aff. liliensterni (MHH coll.), saying "the find to be described here originally originally consisted of a more or less complete skeleton, of which only a few parts had to be recovered" [translated]. He stated the mid caudal centrum "is suitable for identification with Halticosaurus", the tibia is less bowed anteriorly and with a somewhat smaller tuberosity (lateral condyle?), while the metatarsals and phalanges "cannot be distinguished from Halticosaurus liliensterni." The tibia's proximal diameters are 16x9 mm compared to Liliensternus' 95x55 mm, while the caudal is 14 mm long compared to ~53 mm. Thus it is much smaller, even moreso that the co-occuring Pterospondylus, and might be a juvenile or wrongly referred.
References- 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 Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse. Geologische und Palaeontologische Abhandlungen Suppl. 1(1), 1-419.
Jaekel, 1914. Über die Wirbeltierfunde in der oberen Trias von Halberstadt. Paläontologische Zeitschrift. 1, 155-215.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
Huene, 1934. Ein neuer Coelurosaurier in der thüringischen Trias. Paläontologische Zeitschrift. 16(3/4), 145-170.
Kuhn, 1939. Beiträge zur Keuperfauna von Halberstadt. Palaeontologische Zeitschrift. 21, 258-286.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
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.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Rauhut and Hungerbuhler, 2000 (as 1998). A review of European Triassic theropods. Gaia. 15, 75-88.
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)
Middle Norian, Late Triassic
Löwenstein Formation, Trossingen, Germany
?(SMNS 52365) incomplete tooth (FABL ~9 mm) (Sander, 1992)
Comments- SMNS 52365 was found with Plateosaurus specimen 65 in 1942, labeled as Pachysaurus in the collection. Sander (1992) stated it "is very similar to the Frick theropod teeth and probably pertains to Liliensternus." He noted the mesial carina has 9 serrations per 2 mm and the distal carina has 7.5 per 2 mm.
Reference- Sander, 1992. The Norian Plateosaurus bonebeds of central Europe and their taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology. 93, 255-296.

cf. Lilensternus (Sulej, Niedzwiedzki and Bronowicz, 2012)
Middle-Late Norian, Late Triassic
Poręba, Zbaszynek Beds, Poland
Material- (ZPAL V.39/33) partial fused pelvis
Comments- Discovered from 2008-2012, Sulej et al. (2012) referred the pelvis ZPAL V.39.33 to Coelophysoidea indet., and additionally stated the lack of confluence between the supracetabular crest and brevis shelf was shared with Liliensternus. However, Niedzwiedzki et al. (2014) redescribed it as Theropoda indet., finding the condition is also present in Dilophosaurus but not Tawa, Lophostropheus or coelophysids. They also found the notch below the postacetabular process "is most similar to Liliensternus and Lophostropheus, both of which possess deep notches that, like in ZPAL V.39/33, result from the large size of the ischial peduncle and its posterior expansion at its distal tip." Indeed, no differences were noted from Liliensternus except the presence of fusion when the specimen is about 1.2 times smaller.
Sulej et al. also referred partial scapulocoracoid ZPAL V.39/35 to Coelophysoidea, but it was placed in Herrerasauridae by Niedzwiedzki et al. (2014) once described. Similarly, Sulej et al. assigned teeth ZPAL V.39/37, caudal vertebrae ZPAL V.39/38 and pedal unguals ZPAL V.39/36 to Coelophysoidea indet., but Niedzwiedzki et al. reported they "do not preserve any clear neotheropod (or dinosaur or dinosauriform) character states; this material is not described in this article and will be the subject of future study." They are placed in Archosauriformes indet. here.
References- Sulej, Niedzwiedzki and Bronowicz, 2012. A new Late Triassic vertebrate fauna from Poland with turtles, aetosaurs, and coelophysoid dinosaurs. Journal of Vertebrate Paleontology. 32(5), 1033-1041.
Niedzwiedzki, Brusatte, Sulej and Butler, 2014. Basal dinosauriform and theropod dinosaurs from the Mid-Late Norian (Late Triassic) of Poland: Implications for Triassic dinosaur evolution and distribution. Palaeontology. 57(6), 1121-1142.

Notatessaraeraptor Zahner and Brinkmann, 2019
N. frickensis Zahner and Brinkmann, 2019
Rhaetian, Late Triassic
Gruhalde Member of the Klettgau Formation (= Upper Variegated Marls), Switzerland
Holotype- (SMF 06-1; Frickopod) (juevile to subadult) few cervical ribs, first to thirteenth dorsal vertebrae (d2 31 mm, d10 42 mm), several dorsal ribs, gastralia, four sacral vertebrae, sacral ribs, first to fourth caudal vertebrae (c1 28 mm, c4 33 mm), few proximal chevrons, scpulae, coracoid, humeri (~128 mm), radii (97 mm), ulnae, carpals, metacarpal I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1, phalanges II-2 (one proximal), manual ungual II, metacarpals III, phalanx III-1, phalanx III-2, phalanx III-3, metacrapals IV, phalanx IV-1, ilia, pubes, ischia
....(SMF 09-2) incomplete skull (~225 mm), mandible
Referred- ?(SMF 24) tooth (Sander, 1992)
?(SMF 30) incomplete tooth (~13x~6x? mm) (Sander, 1992)
?(SMF 31) tooth (~8x~5x? mm) (Sander, 1992)
?(SMF 32) tooth (~7x~4x? mm) (Sander, 1992)
?(SMF 33) tooth (~13x~7x? mm) (Sander, 1992)
?(SMF coll.) about 11 teeth (5-23 mm) (Sander, 1992)
?(SMF coll.) (~1.5 m) posterior skull, cervical vertebrae, ribs, two ?sacral vertebrae, ilium, tibia, fibula, metatarsal I, phalanx I-1, metatarsal II, phalanx II-1, phalanx II-2, proximal pedal ungual II, metatarsal III, phalanx III-1, proximal phalanx III-2, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV (Anonymous, 2017)
Diagnosis- (after Zahner and Brinkmann, 2019) four exceptionally long but slender premaxillary tooth crowns (as long as anterior maxillary teeth but mesiodistally less wide- ratio 3/1 versus 2.4/1); premaxillary tooth crowns labiolingually flattened, mesially somewhat broader than distally and with fine serrations along their mesial and distal carinae (5 per 1 mm); promaxillary foramen and maxillary fossa present; supratemporal fossa restricted to posterior half of parietal; shallow basisphenoid recess; exit of vagus nerve (X) through a posterior foramen lateral to foramina for hypoglossal nerve (XII); three distinct processes of the articular (medial, dorsolateral and dorsal); markedly low rectangular posterior dorsal neural spines (ratio 2/1); posteriorly increasing height of dorsal neural spines; flattened ventral surfaces and expanded articular faces of sacral centra; deep fossa on lateral surface of second sacral vertebra; proximal caudals with longitudinal fossae on centra and neural arches; prominent anteroproximally located tubercular processes on first four chevrons; pronounced pubic and ischial boots, ischial boot larger than pubic boot.
Comments- The holotype postcrania was discovered in spring 2006, but the skull was only found in 2009. Different portions of the skeleton were described in three theses (Hugi, 2008; Unterrassner, 2009; Zahner, 2014), while Mallison (online 2012) reported on and photographed the specimen as the Frickopod. Oettl-Rieser and Zahner (2018) discussed this as the Frick theropod before Zahner and Brinkmann (2019) officially described and named it. They used a novel phylogenetic analysis to recover Notatessaraeraptor closer to averostrans than coelophysids, Liliensternus and Zupaysaurus, but further than Dracoraptor, Dilophosaurus and Cryolophosaurus. Interestingly, their figure 1i apparently shows the large slab prior to final preparation as photographed by Mallison seven years earlier that exposed the dorsal column. Perhaps it was taken from Hugi's thesis.
Tooth MSF 24 was found in 1985 associated with Plateosaurus skeleton MSF 23, while "about 15 isolated rootless theropod teeth were found during the 1988 excavation" (MSF coll. including MSF 30-33) associated with other Plateosaurus skeletons (1988/1, 1988/2 and/or 1988/3) (Sander, 1992). Sander referred these to cf. Liliensternus based on the high DSDI (~1.08), which he compared to kayentakatae, Lophostropheus and Dilophosaurus, "the teeth of Liliensternus liliensternii" being "generally similar in shape, size, and density of serration." Notatessaraeraptor is very similar to Liliensternus and from the same locality, though its lateral tooth morphology remains unpublished. The teeth are thus provisionally referred to it, pending publication of Zahner's thesis.
In mid May 2017 a theropod skeleton was discovered in the same layer as the isolated teeth (Anonymous, 2017), the published hindlimb photo of which shows is neotheropod. The ilium is similar to Notatessaraeraptor, and all but the hindlimb should be eventually comparable. The article says this may be a new taxon, but it is provisionally listed here given the coelophysoid-grade morphology and proximity to Notatessaraeraptor.
References- Sander, 1992. The Norian Plateosaurus bonebeds of central Europe and their taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology. 93, 255-296.
Hugi, 2008. The axial and appendicular morphology of the first theropod skeleton (Saurischia, Dinosauria) of Switzerland (Late Triassic; Frick, Canton Aargau). Masters thesis, University of Zurich. 161 pp.
Unterrassner, 2009. The anterior appendicular morphology and the stomach content of the first theropod skeleton (Saurischia, Dinosauria) of Switzerland (Late Triassic; Frick, Canton Aargau). Masters thesis, University of Zurich. 136 pp.
Mallison, online 2012. https://dinosaurpalaeo.wordpress.com/2012/05/23/theropod-thursday-19-the-mysterious-frickopod/
Zahner, 2014. Skull morphology of the first theropod skeleton (Saurischia, Dinosauria) from Switzerland (Late Triassic; Frick, Canton Aargau). Masters thesis, University of Zurich. 122 pp.
Anonymous, online 2017. https://www.tagesanzeiger.ch/wissen/natur/Wurde-im-Fricktal-eine-neue-Dinosaurier-Art-entdeckt/story/25194224
Oettl-Rieser and Zahner, 2018. FRICK: Late Triassic basal sauropodomorph and theropod dinosaurs at the Sauriermuseum Frick, Switzerland. In Beck and Joger (eds.). Paleontological Collections of Germany, Austria and Switzerland. Springer International Publishing. 241-248.
Zahner and Brinkmann, 2019. A Triassic averostran-line theropod from Switzerland and the early evolution of dinosaurs. Nature Ecology & Evolution. 3, 1146-1152.

Sarcosaurus Andrews, 1921
= "Liassaurus" Welles, Powell and Pickering vide Pickering, 1995
S. woodi Andrews, 1921
= Magnosaurus woodi (Andrews, 1921) Huene, 1932
= Magnosaurus woodwardi Huene, 1932
= Megalosaurus (Magnosaurus) woodwardi Huene, 1932
= Sarcosaurus andrewsi Huene, 1932
= Megalosaurus andrewsi (Huene, 1932) Waldman, 1974
= "Liassaurus huenei" Welles, Powell and Pickering vide Pickering, 1995
Early Sinemurian, Early Jurassic
Barrow upon Soar, bucklandi Zone, Scunthorpe Mudstone Formation, England
Holotype- (NHMUK R4840/1) (robust) partial posterior dorsal vertebra, partial ilia (248 mm), proximal pubes, ischial fragment, incomplete femur (321.0 mm)
Early Sinemurian, Early Jurassic
Wilmcote, bucklandi Zone, Rugby Limestone Member of the Blue Lias Formation, England
Referred- (WARMS G667-690; intended holotype of "Liassaurus huenei") (7 year old subadult; gracile) partial mid-posterior dorsal vertebra (44.3 mm), five dorsal rib fragments, mid caudal centrum (50.3 mm), ilial fragment, partial pubes, incomplete femora (~317 mm), tibiae (one proximal; ~297 mm), proximal fibula, distal fibula, distal metatarsal II, proximal phalanx II-1, proximal metatarsal II or III, distal metatarsal III, distal metatarsal IV, three fragments (Huene, 1932)
Late Hettangian, Early Jurassic
Wilmcote, angulata zone, Blue Lias Formation, England
(NHMUK R3542; holotype of Magnosaurus woodwardi; holotype of Sarcosaurus andrewsi) tibia (445 mm) (Woodward, 1908)
Diagnosis- (after Andrews, 1921) subpreacetabular notch extremely acute.
(after Ezcurra et al., 2021) ilium with a slightly posteriorly projecting ischiadic peduncle; ilium without laterally exposed ventromedial margin of the brevis fossa except for a short portion of its base (also in Cryolophosaurus); ilium with a poorly transversely expanded brevis fossa (also in other non-coelophysoid neotheropods); femur with dorsolateral trochanter (also in other non-averostran neotheropods); fourth trochanter poorly posteriorly expanded (also present in coelophysoids and early ceratosaurians); femur without an extensor groove; tibia with fibular crest that reaches posterior lateral condyle; tibia with anteroposterior depth versus mediolateral width ratio ≥ 0.6; tibia with anteroposteriorly narrow facet for reception of the ascending process of the astragalus; tibia with an angle between the main axis of the lateral half of the facet for reception of the ascending process of the astragalus and the longitudinal axis of the bone ≥ 25 degrees in anterior view; tibia with a proximally well-extended posteromedial notch on the distal end; tibia with a poorly projected medial malleolus; fibula with a poorly projected and tab-like posterior margin of the proximal end in lateral 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.
Carrano and Sampson (2004) claimed Sarcosaurus is undiagnostic, but Ezcurra et al. (2021) confirmed "the holotype of Sarcosaurus woodi to be diagnosable using a unique combination of character states."
andrewsi- The tibia NHMUK R3542 was originally described by Woodward (1908) as a megalosaurian whose slenderness "and the trochlear shape of the facette for the astragalus, suggest a lighter and more active reptile than the ordinary Megalosaurians", but left it unnamed as "the tibia alone is insufficient for a more exact determination of its affinities." It was then accidentally made the type of two species simultaneously by Huene (1932), Magnosaurus woodwardi and Sarcosaurus andrewsi. 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. Huene (1956) ended up calling it Sarcosaurus andrewsi, making him first reviewer and establishing the correct name. 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. This was formalized by Ezcurra et al. (2021).
"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. The supposed distal pubis is actually a proximal fibula (Ezcurra et al., 2021). Carrano and Sampson (2004) referred to the skeleton 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. As this latter character varies dimorphically in ceratosaur-grade theropods, it is not seen as taxonomically significant. Ezcurra et al. redescribed the specimen and referred it to Sarcosaurus woodi based on "the unique combination of a proportionally short middle-posterior dorsal centrum (length - anterior height ratio < 2, also present in non-coelophysid theropods) and femur with a low fourth trochanter (also present in coelophysids and early ceratosaurian averostrans) and with a dorsolateral trochanter on the proximal end (also present in non-averostran theropods)."
Relationships- Andrews (1921) originally assigned Sarcosaurus 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 1900s. Kurzanov (1989) referred it to his more restricted concept of Megalosauridae (containing Megalosaurus and sinraptorids) without comment. Huene (1932) placed it in Coeluridae based on similarity to Elaphrosaurus (now recognized as a ceratosaur), but in 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. 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. Ezcurra (2012) found Sarcosaurus to be the most basal ceratosaur in a large unpublished analysis. Most recently, Ezcurra et al. (2021) used the Nesbitt dinosauromorph analysis to recover Sarcosaurus closer to Averostra than Dilophosaurus, but less so than Tachiraptor. Correcting some of the scorings moves it to a more basal position closest to Liliensternus and Zupaysaurus.
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.
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. 464 pp.
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(2), 125-136.
Rowe and Gauthier, 1990. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 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.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Thesis. University of Texas at Austin. 553 pp.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Ezcurra, Butler, Maidment, Sansom, Meade and Radley, 2021 (online 2020). A revision of the early neotheropod genus Sarcosaurus from the Early Jurassic (Hettangian-Sinemurian) of central England. Zoological Journal of the Linnean Society. 191(1), 113-149.

Zupaysaurus Arcucci and Coria, 2003
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, 2007) 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, 2007) 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 Averostra); short and square-shaped retroarticular process of the mandible; cnemial crest poorly developed.
Comments- The name "Zupaysaurus" was first announced on the Discovery News television program on 11-12-1999, but was not published until its official description in 2003.
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 (2007). 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 posterolaterally concave distal end; ascending process of astragalus anteriorly 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. The same position was found in Ezcurra's (2012) large unpublished analysis.
References- Arcucci and Coria, 1997. First record of Theropoda (Dinosauria - Saurischia) from the Los Colorados Formation (Upper Triassic, La Rioja, Argentina). XIII Jornadas Argentinas de Paleontologia de Vertebrados, resumenes. Ameghiniana. 34(4), 531.
Arcucci and Coria, 1998. Skull features of a new primitive theropod from Argentina. Journal of Vertebrate Paleontology. 18(3), 24A-25A.
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. 102-104.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Thesis. University of Texas at Austin. 553 pp.
Ezcurra, 2007 (online 2006). The cranial anatomy of the coelophysoid theropod Zupaysaurus rougieri from the Upper Triassic of Argentina. Historical Biology. 19(2), 185-202.
Ezcurra and Novas, 2007 (online 2006). Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
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.
Paulina-Carabajal, 2009. El neurocráneo de los dinosaurios Theropoda de la Argentina: Osteología y sus implicancias filogenéticas. PhD Thesis, Universidad Nacional de La Plata. 554 pp.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Paulina Carabajal, Ezcurra and Novas, 2015. New information on the braincase and endocranial morphology of the Late Triassic theropod Zupaysaurus rougieri using CT scans. XXIX Jornadas Argentinas de Paleontología de Vertebrados, resumenes. Ameghiniana. 52(4) suplemento, 32.

unnamed clade (Cryolophosaurus ellioti + Dilophosaurus wetherilli + Passer domesticus)
Diagnosis- (suggested) antorbital fossa extends on to lateral surface of nasal; lacrimal horn; anteroposteriorly shortened orbit; highest point of axial neural spine near posterior edge; cervical ribs with pneumatic fossae (also in right tenth rib of Lucianovenator).

unnamed dilophosaur (Milner and Lockley, 2006)
Hettangian, Early Jurassic
Freeman Quarry SGDS14V, Whitmore Point Member of the Moenave Formation, Utah, US
Material- (SGDS 768) ~sixth dorsal vertebra (41.38 mm)
Comments- This was discovered on January 4 2004. Kirkland and Milner (2005) first mention "Bones and teeth of theropods large enough to produce Eubrontes tracks" from the Whitmore Point Member, while Milner and Lockley (2006) state "A mid-dorsal vertebra of an unknown ceratosaur could also belong to Megapnosaurus (Fig. 7H). This specimen is currently under preparation and still requires detailed study." They figure it in posterior view as "mid-dorsal vertebra from a coelophysoid theropod dinosaur." Milner et al. (2012) figure the vertebra in multiple views as "Theropod cranial thoracic (anterior dorsal) vertebra (SGDS 768)", and it was later described in detail by Marsh et al. (2021). The latter study used Nesbitt's dinosauromorph analysis to recover it as a member of the Dilophosaurus + Averostra clade. Comparison to Dilophosaurus suggests it is around the sixth dorsal based on parapophysis position, neural spine shape in dorsal view and centrum shape in ventral view, but it differs in having more transversely flared centrum ends and retaining a ventral keel. Notably this eliminates the supposed difference of lacking a pleurocoel, since these are only present in the first four dorsals of Dilophosaurus.
References- Kirkland and Milner, 2005. The case for theropod dinosaurs exploiting fish as a major food resource during the Early Jurassic. Tracking Dinosaur Origins: The Triassic/Jurassic Terrestrial Transition Abstracts Volume. 9-10.
Milner and Lockley, 2006. History, geology, and paleontology: St. George Dinosaur Discovery Site at Johnson Farm, Utah. In Reynolds (ed.). Making Tracks Across the Southwest, Abstracts from the 2006 Desert Symposium. 35-48.
Milner, Birthisel, Kirkland, Breithaupt, Matthews, Lockley, Santucci, Gibson, DeBlieux, Hurlbut, Harris and Olsen, 2012. Tracking Early Jurassic dinosaurs across southwestern Utah and the Triassic-Jurassic transition. Nevada State Museum Paleontological Papers. 1, 1-107.
Marsh, Milner, Harris, De Blieux and Kirkland, 2021. A non-averostran neotheropod vertebra (Dinosauria: Theropoda) from the earliest JurassicWhitmore Point Member (Moenave Formation) in southwestern Utah. Journal of Vertebrate Paleontology. e1897604.

undescribed possible dilophosaur (Kutty, Chatterjee, Galton and Upchurch, 2007)
Sinemurian, Early Jurassic
Upper Dharmaram Formation, India
Material- teeth, limb fragments
Comments- Kutty et al. state this is similar to Dilophosaurus.
Reference- Kutty, Chatterjee, Galton and Upchurch, 2007. Basal sauropodomorphs (Dinosauria: Saurischia) from the Lower Jurassic of India: Their anatomy and relationships. Journal of Paleontology. 81, 1218-1240.

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
Referred- material (Smith et al., 2012)
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 Averostra. 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- Monastersky, 1993. From Antarctica: The Elvis of dinosaurs. Science News. 144(17), 261-261.
Holmes, 1993. [title]. Prehistoric Times. [issue, pp]
Hammer and Hickerson, 1994. A crested theropod dinosaur from Antarctica. Science. 264, 828-830.
Hammer, Hickerson and Slaughter, 1994. A dinosaur assemblage from the Transantarctic Mountains. Antarctic Journal. 29(5), 31-33.
Smith, 2005. Osteology of Cryolophosaurus ellioti (Dinosauria, Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Masters Thesis, University of Iowa. 384 pp.
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.
Smith, Hellert, Mathews, Hammer and Makovicky, 2012. New dinosaurs from the Early Jurassic Hanson Formation of Antarctica, and patterns of phylogemetic diversity in Early Jurassic sauropodomorphs. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 175.

Dilophosauridae Madsen and Welles, 2000
= "Dilophosauridae" Charig and Milner, 1990
= Dilophosaurinae Madsen and Welles, 2000
Definition- (Dilophosaurus wetherilli <- Coelophysis bauri, Ceratosaurus nasicornis, Allosaurus fragilis) (Holtz, 2012)
Other definitions- (Dilophosaurus wetherilli <- Coelophysis bauri, Ceratosaurus nasicornis, Passer domesticus) (Hendrickx, Hartman and Mateus, 2015)
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 Averostra. Smith et al. (2007) later found a clade containing "Dilophosaurus" sinensis (= Sinosaurus), 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, 2007) or sister to dilophosaurids plus more derived theropods (Ezcurra, 2012). Smith et al.'s dilophosaur clade was also sister to Averostra. 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.
References- Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 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. 127-140.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised osteology. Miscellaneous Publication 00-2 Utah Geological Survey. 80 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.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Thesis. 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, 2007 (online 2006). Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
Smith and Makovicky, 2007. Early theropod evolution and paraphyly of the Coelophysoidea. Journal of Vertebrate Paleontology. 27(3), 150A.
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.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Holtz, 2012. Theropods. In Brett-Surman, Holtz and Farlow (eds.). The Complete Dinosaur 2nd Edition. Indiana University Press. 346-378.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Dilophosaurus Welles, 1970
D. wetherilli (Welles, 1954) Welles, 1970
= Megalosaurus wetherilli Welles, 1954
= Dilophosaurus "breedorum" Welles, 1995 vide Welles and Pickering, 1995
Sinemurian-Pliensbachian, Early Jurassic
Moa Ave 1 UCMP V4214, Silty Facies Member of the 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? (lost), intermedium? (lost), distal carpal I, distal carpal II? (lost), distal carpal III? (lost), distal carpal IV? (lost), 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)
Paratype- (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
Sinemurian-Pliensbachian, Early Jurassic
Rock Head MNA 219-0, Silty Facies Member of the Kayenta Formation, Arizona, US
Referred- (MNA.V.97) tooth (Gay, 2001)
....(MNA.V.101) proximal tibia (Gay, 2001)
?...(MNA.V.116) manual phalanx III-1 (48 mm) (Gay, 2001)
....(MNA.V.131) pedal phalanx III-1 (120 mm) (Gay, 2001)
....(MNA.V.135) incomplete cervical vertebra (83 mm) (Gay, 2001)
?...(MNA.V.141) manual phalanx III-2 (60 mm) (Gay, 2001)
?...(MNA.V.142) partial rib (Gay, 2001)
....(MNA.V.154) proximal pubis (Gay, 2001)
....(MNA.V.160) proximal femur (~545 mm) (Gay, 2001)
....(MNA.V.161) distal femur (Gay, 2001)
....(MNA.V.176) dorsal centrum (31 mm) (Gay, 2001)
....(MNA.V.177) partial proximal caudal vertebra (85 mm), partial neural spine (Gay, 2001)
....(MNA.V.248) proximal tibia (Gay, 2001)
....(MNA.V.530) proximal fibula (Gay, 2001)
....(MNA.V.539) proximal fibula (Gay, 2001)
Sinemurian-Pliensbachian, Early Jurassic
Gold Spring E (= Blue Valley) MNA 356-0, Silty Facies Member of the Kayenta Formation, Arizona, US
(MNA.V.3145) distal femur (~574 mm) (Gay, 2005)
Late Pliensbachian, Early Jurassic
Dilophosaurus Quarry TMM 43646, Silty Facies Member of the Kayenta Formation, Arizona, US
(TMM 43646-1) (~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)
Sinemurian-Pliensbachian, Early Jurassic
TMM 43662, Silty Facies Member of the Kayenta Formation, Arizona, US
(TMM 43662-2) (gracile adult) proximal femur (Tykoski, 2005)
Sinemurian-Pliensbachian, Early Jurassic
Paiute North 1 TMM 43691, Silty Facies Member of the Kayenta Formation, Arizona, US
(TMM 43691-1) ilium (Marsh and Rowe, 2020)
Sinemurian-Pliensbachian, Early Jurassic
Jon's Theropod TMM 47006, Silty Facies Member of the Kayenta Formation, Arizona, US
(TMM 47006-1) braincase, presacral column (Marsh and Rowe, 2020)
Sinemurian-Pliensbachian, Early Jurassic
Moa Ave 2 UCMP V6468, Silty Facies Member of the Kayenta Formation, Arizona, US
(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)
Sinemurian-Pliensbachian, Early Jurassic
Pumpkin Patch UCMP V6899, Silty Facies Member of the Kayenta Formation, Arizona, US
(UCMP 130083) scapulocoracoid fragments (UCMP online)
Sinemurian-Pliensbachian, Early Jurassic
Rock Head 2 UCMP V82303, Silty Facies Member of the Kayenta Formation, Arizona, US
(UCMP 130053) vertebral fragments (Clark and Fastovsky, 1986)
Sinemurian-Pliensbachian, Early Jurassic
Willow Springs 6 UCMP V82313, Silty Facies Member of the Kayenta Formation, Arizona, US
(UCMP 130069) ischium (UCMP online)
(UCMP 130070) vertebrae, proximal femur, limb fragment, phalanx (UCMP online)
Sinemurian-Pliensbachian, Early Jurassic
ARCH 71v, Sandy Facies Member of the Kayenta Formation, Utah, US
?(ARCH 4012) vertebral fragments, limb fragments, metapodial fragments (Madsen, Kirkland, DeBlieux, Santucci, Inkenbrandt and Tweet, 2012)
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.
(after Marsh, 2015) preorbital boss on lacrimal; posterior centrodiapophyseal lamina bifurcates and reunites down the neck so that the single posterior centrodiapophyseal lamina present on the third cervical is not homologous to that found on the eighth cervical; coracoid has a ventral tubercle; scalloped obturator process; notch on posteromedial corner of distal tibia.
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.
Marsh and Rowe (2020) reidentified several of the elements described by Gay (2001)- proximal pubis MNA.V.101 is a proximal tibia; MNA.V.135 is a cervical vertebra not a dorsal vertebra; MNA.V.177 is a caudal vertebrae instead of a dorsal; pubis MNA.V.248 is a proximal tibia, though that leaves a mystery where the rest of the element went as Gay measures the whole thing as 570 mm; MNA.V.530 and 539 are proximal instead of distal fibulae. MNA.V.116, 141 and 142 are not mentioned, so may be cf. Dilophosaurus or Archosauria indet.. arsh and Rowe also reidenfified supposed proximal femur MNA.V.3145 (Gay, 2005) as a distal femur.
Not Dilophosaurus- Gay (2001) described numerous specimens from the Rock Head locality as Dilophosaurus, but Marsh and Welles (2020) removed femur MNA.V.109 as ornithischian, and distal ?fibula MNA.V.102, dentary fragments MNA.V.111, ?ischial fragment MNA.V.122, caudal centrum MNA.V.138 and rib fragment MNA.V.247 as "incongruent in size with the Dilophosaurus remains and attributable to Archosauria."
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 "Syntarsus" 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. Marsh (2012) recently also only recognized a single species in his restudy of the taxon.
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. in Fraser and Sues (eds.). The Beginning of the Age of the Dinosaurs: Faunal change across the Triassic-Jurassic boundary. 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 1999. https://web.archive.org/web/20200712043408/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. Thesis, University of Bristol. 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. Mesa Southwest Museum Bulletin. 8, 19-23.
Gay, 2005. Sexual dimorphism in the Early Jurassic theropod dinosaur 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 Thesis. 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 (online 2007). 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.
Madsen, Kirkland, DeBlieux, Santucci, Inkenbrandt and Tweet, 2012. Paleontological resources inventory and monitoring, Arches National Park, Utah. Utah Geological Survey Contract Deliverable. Cooperative Agreement #H230097080, 119 pp.
Marsh, 2015. A comprehensive study of Dilophosaurus wetherilli: Anatomy, taxonomy, and evolutionary relationships of the first large-bodied theropod in North America. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 175.
Senter, 2015. Range of motion in the forelimb of the theropod dinosaur Dilophosaurus wetherilli. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 212.
Senter and Juengst, 2016. Record-breaking pain: The largest number and variety of forelimb bone maladies in a theropod dinosaur. PLoS ONE. 11(2), e0149140.
Marsh and Rowe, 2020. A comprehensive anatomical and phylogenetic evaluation of Dilophosaurus wetherilli (Dinosauria, Theropoda) with descriptions of new specimens from the Kayenta Formation of northern Arizona. Journal of Paleontology. 94(Memoir 78), 103 pp.

Dracovenator Yates, 2005
D. regenti Yates, 2005
Pliensbachian, Early Jurassic
Upper Drumbo Farm, Upper Elliot Formation or Clarens Formation, South Africa
Holotype- (BP/1/5243) premaxillae, maxillary fragment, two dentary fragments, partial surangular, partial angular, partial prearticular, articular, teeth
Diagnosis- (after Yates, 2005) 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.
Comments- Rauhut and Lopez Arbarello (2008) stated "Recent fieldwork indicates, however, that this specimen might be derived from the basal part of the overlying Clarens Formation (OR, pers. obs. 2008)."
Yates (2005) tentatively referred snout BP/1/5278 to Dracoventor, but Ezcurra (2012) found it to be a non-coelophysid coelophysoid sensu stricto in a large unpublished analysis, while Dracovenator was still a dilophosaurid. Indeed, Wang et al. (2017) noted "None of the autapomorphies of Dracovenator can be observed on BP/1/5278, so this specimen is included as a separate OTU in this analysis to test its affinities", where it emerged as a coelophysid while Dracovenator was sister to Dilophosaurus plus averostrans.
References- 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.
Rauhut and López-Arbarello, 2008. Archosaur evolution during the Jurassic: A southern perspective. Revista de la Asociación Geológica Argentina. 63(4), 557-585.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 91.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.

unnamed clade (Tachiraptor admirabilis + Passer domesticus)
Diagnosis- (suggested) medial malleolus of tibia angled from the shaft (also in Zupaysaurus); distal tibia anteroposterior depth <60% of transverse width.

Tachiraptor Langer, Rincon, Ramezani, Solorzano and Rauhut, 2014a
T. admirabilis Langer, Rincon, Ramezani, Solorzano and Rauhut, 2014a
Hettangian, Early Jurassic
La Quinta Formation, Venezuela
Holotype- (IVIC-P-2867) (~1.5 m) incomplete tibia (~250 mm)
Paratype- ?...(IVIC-P-2868) proximal ischium
Diagnosis- (after Langer et al., 2014a) posterolateral corner of fibular condyle forms sharp angle in proximal view and extends slightly more posteriorly than the medial condyle; (combination of) distal articulation of tibia more than 50% broader transversely than anteroposteriorly; astragalar buttress occupies between one-third and one-quarter of anteroposterior depth of distal surface of bone, extending obliquely across the anterior surface of distal part of the tibia at an angle of approximately 35 degrees to the distal margin, and flexing proximally at the lateral 20% of the transverse width of the distal shaft; line connecting the outer and inner tibial malleoli in anterior view forms angle of ~80 degrees to long axis of bone.
Comments- Langer et al. (2014a,b) added this taxon to Smith et al.'s basal theropod dataset and found it to be sister to Averostra. As the original matrix is heavily un/miscoded, an assignment to basal Ceratosauria or Tetanurae may not be unlikely.
References- Langer, Rincon, Ramezani, Solorzano and Rauhut, 2014a. New dinosaur (Theropoda, stem-Averostra) from the Earliest Jurassic of the La Quinta Formation, Venezuelan Andes. Royal Society Open Science. 1, 140184.
Langer, Rincon, Ramezani, Solorzano and Rauhut, 2014b. New theropod material from the Triassic-Jurassic boundary of the Venezuelan Andes. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 165.