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 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 Pal�ontologische 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). Pal�ontologische 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). Pal�ontologische
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 (Sarig�l, 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, Sarig�l (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- Sarig�l, 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.
Sarig�l, 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.
Sarig�l, 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.
Sarig�l, 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 neotheropod (Owen, 1859)
Hettangian-Early Sinemurian, Early Jurassic
Blue Lias Formation, England
Material- (NHMUK 39496; holotype of Scelidosaurus harrisonii) distal
femur (~640 mm), proximal tibia, proximal fibula (lost)
Diagnosis- (suggested) combination of bulbous fibular crest on tibia
and very shallow extensor groove on distal femur.
Comments- Owen received the partial hindlimbs NHMUK 39496 and GSM 109560
in 1858 and used them as the basis for his dinosaur genus Scelidosaurus
in an encyclopedia entry the following year. While this is often claimed to
be an nomen nudum (e.g. Newman, 1968), genus names published before 1931 do
not require species names or illustrations to be valid (ICZN Article 12). Owen
later (1861) gave his taxon the species name harrisonii and described
it in detail, referring the ungual GSM 109561, a partial postcranium in the
Lyme Regis Museum, and the skull of NHMUK R1111. Lydekker (1888) made NHMUK 39496
the type specimen, though the basal thyreophoran NHMUK R1111 (whose postcranium
was soon found and described in 1862) formed the basis for peoples' ideas of
Scelidosaurus. Newman (1968) believed NHMUK 39496 and GSM 109560 to be
megalosaurids, and possibly GSM 109561 as well. However, they were only compared
to Megalosaurus among theropods, making this familial assignment in need
of verification. As the name Scelidosaurus had been associated with the
thyreophoran, Charig and Newman (1994) petitioned the ICZN to recognize NHMUK
R1111 as the lectotype, which was accepted in 1994 as Opinion 1788. Welles and
Powell studied the theropod material in 1974 for their unpublished European
theropod paper, intending to name it Merosaurus newmani.
This was first found in publically available print in 1995 when
Pickering credited the name to Welles, Powell and Pickering in an
unpublished bibliographic manuscript. In that same year, Pickering
printed a packet with a full description of the taxon. 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.
Pickering intends GSM 109560 to be the type, and referred NHMUK 39496
and GSM 109561. He considered it a probable ceratosaur sensu lato
metataxon. Pickering will describe it in his in progress work Mutanda
Dinosaurologica. Carrano and Sampson (2004) stated NHMUK 29496 probably
belongs to a basal tetanurine without explanation, but viewed it as
indeterminate. Naish and Martill (2007) referred all three specimens to
Tetanurae without comment. Most recently, Benson (2009, 2010)
redescribed NHMUK 39496 and GSM 109560. He found NHMUK 39496 to be a
coelophysoid when entered in his matrix, but noted that non-tetanurines
were poorly sampled so this may not mean much. Benson considered it to
be Theropoda incertae sedis, and the other two specimens to be
indeterminate theropods. Carrano et al. (2012) noted it may be
tetanurine, but they were unsure. When added to their matrix, NHMUK
39496 emerges in a polytomy with Cryolophosaurus, "Dilophosaurus"
sinensis, Ceratosauria and Tetanurae.
NHMUK 39496 consists of a distal femur, proximal tibia and a proximal fibula
which has been lost subsequent to Newman's description. Both Pickering and Benson
proposed the lack of a deep extensor groove as a non-tetanurine character, but
this is true in some basal tetanurines (Chuandongocoelurus), megalosauroids
(Dubreuillosaurus, Eustreptospondylus, "Brontoraptor")
and most coelurosaurs. Contra Pickering, the popliteal notch is concave and
the ectocondyle elliptical and posterolaterally directed even in tetanurines
like Megalosaurus and Eustreptospondylus. Benson claims the fibular
crest which extends to the lateral condyle is a non-tetanurine character, but
this is also found in Afrovenator, Megalosaurus and Gasosaurus.
One feature of the tibia might suggest this specimen is a tetanurine- the fibular
crest is bulbous as in Piatnitzkysaurus, Megalosaurus and Sinraptor.
However, including NHMUK 39496 in my saurischian supermatrix results in equally
parsimonious trees where it is in Coelophysoidea or non-avetheropod Tetanurae.
Morphologies in the preserved areas are not consistantly variable between these
groups, making further identification difficult. Thus it is here assigned to Neotheropoda incertae sedis.
References- Owen, 1859. Palaeontology. Encyclopaedia Britannica, Edition
8. 17, 91-176.
Owen, 1861. Monograph of the fossil Reptilia of the Liassic formations. Part
I. A monograph of the fossil dinosaur (Scelidosaurus harrisonii Owen)
of the Lower Lias. Palaeontolographical Society Monographs. 13, 1-14.
Owen, 1862. Monographs on the British Fossil Reptilia from the Oolitic Formations.
Part second, containing Scelidosaurus harrisonii and Pliosaurus grandis.
Palaeontolographical Society Monographs. 1-16.
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, Squamata, Rhynchocephalia, and Proterosauria.
British Museum of Natural History, London. 309 pp.
Newman, 1968. The Jurassic dinosaur Scelidosaurus harrisoni Owen. Palaeontology.
11, 40-43.
Charig and Newman, 1992. Scelidosaurus harrisonii Owen, 1861 (Reptilia,
Ornithischia): Proposed replacement in inappropriate lectotype. Bulletin of
Zoological Nomenclature. 49, 280-283.
ICZN, 1994. Opinion 1788. Scelidosaurus harrisonii Owen, 1861 (Reptilia,
Ornithischia): Lectotype replaced. Bulletin of Zoological Nomenclature. 51(3),
288.
Pickering, 1995a. Jurassic Park: Unauthorized Jewish Fractals in Philopatry.
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Pickering, 1995b. An extract from: Archosauromorpha: Cladistics and osteologies.
A Fractal Scaling in Dinosaurology Project. 11 pp.
Olshevsky, DML 1999. https://web.archive.org/web/20210309141613/http://dml.cmnh.org/1999Dec/msg00193.html
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.
Naish and Martill, 2007. Dinosaurs of Great Britain and the role of the Geological
Society of London in their discovery: Basal Dinosauria and Saurischia. Journal
of the Geological Society. 164, 493-510.
Benson, 2009. The taxonomy, systematics and evolution of the British theropod
dinosaur Megalosaurus. PhD thesis. University of Cambridge. [pp].
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.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Pickering, in prep. Mutanda Dinosaurologica.
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 m�sozo�ques de Normandie.
Ecosyst�mes continentaux m�sozoiques de Normandie (Livret-guide). Universit� 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. Pal�ontologie des vert�br�s
au Grand-Duch� de Luxembourg: D�couvertes r�centes et travaux
en cours. Archives de l’Institut Grand-Ducal de Luxembourg, Section des
Sciences Naturelles, Physiques et Math�matiques, Nouvelle S�rie.
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 L�wenstein 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
w�rttembergischen 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) dorsal centra, proximal femur, partial astragalus,
metatarsal fragments, phalanges
Comments- NMMNH P4569 was originally seen as a coelophysoid by Lucas
et al. (1985) and Hunt and Lucas (1989), though 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. It may therefore be a coelophysoid, as originally believed.
Nesbitt et al. further noted the fragmentary remains referred to "Comanchesaurus"
are indeterminate.
Although the name "Comanchesaurus kuesi" was originally used in thesis,
and thus not available for use in this website, it was later published by Nesbitt
et al. (2007).
References- Lucas, Hunt and Bennett, 1985. Triassic vertebrates from
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.
Dolichosuchus Huene, 1932
D. cristatus Huene, 1932
Early Norian, Late Triassic
Kaltental, L�wenstein 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)
Diagnosis- Provisionally indeterminate relative to Coelophysis bauri.
Comments- This specimen was originally briefly described and illustrated
as Procompsognathidae gen. et sp. indet. by Parrish and Carpenter (1986). It
was then described and named "Revueltoraptor lucasi" by Hunt (1994)
in his unpublished thesis, and called herrerasaurid A by Hunt et al. (1998).
It was finally officially named and described as the coelophysoid Gojirasaurus
quayi by Carpenter (1997). However, Nesbitt et al. (2005) find the holotype
to be chimaeric, which was elaborated on in their 2007 publication. They found
the dorsal vertebrae were referrable to Shuvosaurus and the pubis and
tibia referrable to a coelophysoid (contra the 2005 abstract where the dinosaurian
elements were relegated to Saurischia indet.). They could not assign the other
material (teeth, dorsal ribs, gastralia, chevron) or the several referred specimens
noted by Hunt (1994) (NMMNH P4666, P16607, P16656, P16946, P17134, P17154, P17258,
UMMP 7274) to a particular taxon of archosaur. The reassignment of the dorsal
material makes the sole apomorphy identified by Rauhut invalid (mid/posterior
dorsal vertebrae with taller neural spines than other coelophysoids), and Gojirasaurus
is presently indeterminate relative to Coelophysis except for the greater
robustness of the tibia (Nesbitt et al., 2007). This may be size-related.
Although the name "Revueltoraptor lucasi" was originally used in thesis,
and thus not available for use in this website, it was later published by Nesbitt
et al. (2007). The name was first noted publically on the Dinosaur Mailing List
in 2000 by Gay, who noted several elements at the NMMNH were labeled as "Revueltoraptor".
These were specimens referred to the taxon by Hunt (1994), and are not identifiable
as Gojirasaurus.
Rauhut (2003) suggested Gojirasaurus may be synonymous with Shuvosaurus,
as a large Shuvosaurus premaxilla was found in the same deposits (identified
as Reptilia indet. by Parrish and Carpenter, 1986), and the tooth associated
with the Gojirasaurus holotype may not belong to it. However, Nesbitt
and Norell (2006) have 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.
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. Pal�ontologie und Phylogenie der Niederen Tetrapoden [Paleontology
and Phylogeny of the Lower Tetrapods]. VEB Gustav Fischer Verlang, Jena. 1-716.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica, 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 Lowenstein 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 Europ�ischen Triasformation
mit ber�cksichtigung der Ausseurop�ischen vorkommnisse [The dinosaurs
of the European Triassic formations with consideration of occurrences outside
Europe]. Geologische und Palaeontologische Abhandlungen Suppl. 1(1), 1-419.
Huene, 1909. Skizze zu einer Systematik und Stammesgeschichte der Dinosaurier
[Sketch of the systematics and origins of the dinosaurs]. Centralblatt f�r
Mineralogie, Geologie und Pal�ontologie. 1909, 12-22.
Zittel, 1911. Grundz�ge der Pal�ontologie (Pal�ozoologie). II.
Abteilung. Vertebrata [Fundamentals of Paleontology (Paleozoology). Section
II. Vertebrata]. Druck und Verlag von R. Oldenbourg, M�nchen. 1-598.
Huene, 1914. Das nat�rliche System der Saurischia [The systematics of the
Saurischia]. Centralblatt f�r Mineralogie, Geologie und Pal�ontologie.
1914, 154-158.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien
[Phylogenetic results of some investigations of Triassic reptiles]. Zeitschrift
f�r Induktive Abstammungsund Vererbungslehre. 24, 159-163.
Huene, 1921. Coelurosaurier-Reste aus dem obersten Keuper von Halberstadt. Centralblatt
f�r Mineralogie, Geologie und Pal�ontologie. 1921(10), 315-320.
Bock, 1952. Triassic reptilian tracks and trends of locomotive evolution. Journal
of Paleontology. 26(3), 395-433.
Huene, 1956. Pal�ontologie und Phylogenie der Niederen Tetrapoden [Paleontology
and Phylogeny of the Lower Tetrapods]. VEB Gustav Fischer Verlang, Jena. 1-716.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press,
Chicago. 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.
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277-342.
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Viohl, 1992. No evidence of Triassic birds. Archaeopteryx. 10, 77-79.
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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 g�richi
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 th�ringischen
Trias. Pal�ontologische 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. Beitr�ge zur geschichte der Archosaurier
[Contribution to the history of the archosaurs]. Geologie und Pal�ontologie
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
[Phylogenetic relationships of ceratosaurian theropod dinosaurs]. 28(3-4), 401.
Novas, 1992. La evolucion de los dinosaurios carnivoros [The evolution of carnivorous
dinosaurs]. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico:
Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez",
Cuenca, Argentina. 126-163.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Charig and Milner, 1997. Baryonyx walkeri, a fish-eating dinosaur from
the Wealden of Surrey. Bulletin of the Natural History Museum of London (Geology).
53, 11-70.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. 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 Societ� 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 Pal�ontologische 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 Pal�obiologie. 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. �ber die Wirbeltierfunde in der oberen
Trias von Halberstadt.
Pal�ontologische 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 R�thien 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 f�r Geologie und Pal�ontologie,
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. (Tri�sico Superior) I parte. Opera Lilloana. 22,
1-183.
Novas, 1989. Los dinosaurios carn�voros 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.
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[Contribution to the history of the archosaurs]. Geologie und Pal�ontologie
Abhandlungen. 13(7), 1-56.
Huene, 1914b. Das nat�rliche System der Saurischia [The systematics of
the Saurischia]. Centralblatt f�r Mineralogie, Geologie und Pal�ontologie.
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Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien
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f�r Induktive Abstammungsund Vererbungslehre. 24, 159-163.
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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 �bersicht �ber die Saurischia und ihre nat�rlichen
Zusammenh�nge.
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Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of
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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.
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Steel, 1970. Part 14. Saurischia. Handbuch der Pal�oherpetologie/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 [On the evolution of carnivorous
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Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
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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 Ricql�s 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). Pal�ontologische 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 Ricql�s, Padian and Horner, 2003. On the bone histology of some
Triassic pseudosuchian archosaurs and related taxa. Annales de
Pal�ontologie. 89(2), 67-101.
Padian, Horner and De Ricql�s, 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, M�ller, 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 Mus�um 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. D�couvertes r�centes de Dinosaures au Maroc. In Taquet
and Sudre (eds.). Les Dinosaures de la Chine � la France. Mus�e
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 f�r Geologie und
Pal�ontologie, 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 Lowenstein 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 schw�bischen Dinosaurier. Jahreshefte
des Vereins f�r Vaterl�ndische Naturkunde in W�rttemberg. 68,
66-67.
Fraas, 1913. Die neuesten Dinosaurierfunde in der schwabischen Trias. Naturwissenschaften.
45, 1097-1100.
Huene, 1921. Neue Pseudosuchier und Coelurosaurier aus dem w�rttembergischen
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 f�r Geologie und
Pal�ontologie, 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). Pal�ontologische 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.
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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 Eichst�tt, Eichst�tt. 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 f�r Geologie und
Pal�ontologie 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, Th�ringen, 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 Europ�ischen Triasformation mit ber�cksichtigung
der Ausseurop�ischen vorkommnisse. Geologische und Palaeontologische Abhandlungen
Suppl. 1(1), 1-419.
Jaekel, 1914. �ber die Wirbeltierfunde in der oberen
Trias von Halberstadt.
Pal�ontologische 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 th�ringischen
Trias. Pal�ontologische Zeitschrift.
16(3/4), 145-170.
Kuhn, 1939. Beitr�ge 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
Lowenstein 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. Pal�ontologie 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 neurocr�neo de los dinosaurios Theropoda
de la Argentina: Osteolog�a y sus implicancias filogen�ticas.
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 Paleontolog�a 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 L�pez-Arbarello, 2008. Archosaur evolution during the Jurassic:
A southern perspective. Revista de la Asociaci�n Geol�gica 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.