Coelophysoidea Nopcsa, 1928 sensu Holtz,
1994
Definition- (Coelophysis bauri <- Carnotaurus sastrei)
(modified from Sereno, 1998)
Other definitions- (Coelophysis bauri <- Ceratosaurus nasicornis)
(modified from Padian et al., 1999)
(Coelophysis bauri <- Ceratosaurus nasicornis, Carnotaurus
sastrei, Passer domesticus) (Sereno, in press)
= Podokesauroidea Huene, 1914 sensu Madsen and Welles, 2000
= Ceratosauria sensu Sereno, 1998
Definition- (Coelophysis bauri <- Passer domesticus) (modified)
= Coelophysoidea sensu Padian et al., 1999
Definition- (Coelophysis bauri <- Ceratosaurus nasicornis)
(modified)
= Coelophysoidea sensu Sereno, in press
(Coelophysis bauri <- Ceratosaurus nasicornis, Carnotaurus
sastrei, Passer domesticus)
Diagnosis- (after Tykoski, 2005) supracetabular crest flares lateroventrally
to hide dorsal acetabulum in lateral view; anterior trochanter conical.
Comments- This clade may be part of Ceratosauria (Tykoski, 2005). See
Comments under Ceratosauria.
History of use and Podokesauroidea- The idea of grouping Dilophosaurus
with coelophysoids has a long history, but prior to 1994 was done under two
older names- Podokesauridae (Russell, 1984; Carroll, 1988) and Coelophysidae
(Paul, 1988; Novas, 1991; Novas, 1992). The clade was left unnamed by Rowe (1989),
where it was first supported cladistically. Welles (1984) and some other authors
had recognized the relationship between Dilophosaurus and Liliensternus,
generally placing them both in Halticosauridae, but had only vague ideas about
their relationship with smaller coelophysids. Holtz (1994) was the first to
use the name Coelophysoidea for a Dilophosaurus + Coelophysidae clade,
which was soon defined to be stem-based (Sereno, 1998) and thus included Coelophysis
and any taxa more closely related to it than to neoceratosaurs. The term has
been used in this way ever since. Madsen and Welles (2000) correctly noted that
Podokesauroidea has priority over Coelophysoidea according to the ICZN, since
it was named fourteen years earlier. Yet all other workers ignore this, generally
because they value Phylocode rules and Podokesauroidea has not been phylogenetically
defined, or they misread the ICZN to demand family-level eponyms be diagnosable
(it actually only says they must be valid). Coelophysoidea is only used on this
website because of its near universal usage since 1994.
Is Dilophosaurus a coelophysoid?- An additional phylogenetic issue is
whether Dilophosaurus and related taxa (dilophosaurids as used here)
are coelophysoids, or more closely related to ceratosaurs and tetanurines. Traditional
phylogenies placed them in Coelophysoidea (see above), and this has been found
in many recent analyses as well (Wilson et al., 2003; Holtz et al., 2004; Tykoski
and Rowe, 2004; Tykoski, 2005; Ezcurra and Novas, 2006). Yet a few recent analyses
have instead recovered Dilophosaurus as more closely related to ceratosaurs
and tetanurines (Rauhut, 2003; Yates, 2005; Smith et al., 2007). Smith et al.
recovered the following unambiguous synapomorphies for a dilophosaurid+ceratosaur+tetanurine
clade- dorsoventrally elongate orbit; maxillae that are orientated subparallel
to each other in dorsal aspect; reduced maxillary tooth count; tongue-like process
of the parietals overlapping the supraoccipital knob; relatively deep surangular.
Yet Rauhut found Dilophosaurus to be equally well supported as a coelophysoid
when Shuvosaurus was (correctly) excluded, Yates found dilophosaurids
were coelophysoids with only one added step, and Smith et al. found dilophosaurids
were coelophysoids with only six added steps. Similarly, Ezcurra and Novas found
equal bootstrap support for either option. Yet Tykoski (2005) found it took
twenty additional steps to place Dilophosaurus closer to ceratosaurs
and tetanurines. Perhaps importantly, none of the analyses supporting coelophysoid
dilophosaurids have included Cryolophosaurus, Dracovenator or
"Dilophosaurus" sinensis, whereas Smith et al. included all
three and Yates includes Dracovenator. Also relevent is that Tykoski
found Dilophosaurus to have been miscoded in prior analyses, due to a
reliance on subadult specimens. Whether this was corrected for in later analyses
has yet to be examined. If dilophosaurids are not coelophysoids, the coelophysoid
identity of other taxa such as Sarcosaurus, Halticosaurus, Lophostropheus
and Zupaysaurus comes into question as well. Indeed, both Smith et al.
and Yates found Zupaysaurus to be outside of Coelophysoidea in their
analyses.
Ex-coelophysoids- Several taxa have been included in authors' equivalents
of Coelophysoidea in the past, but do not appear to belong there. Avipes,
Lukousaurus, Saltopus and Velocipes were all included in
Carroll's Podokesauridae, as was common in the pre-cladistic age. None have
ever been placed in Coelophysoidea based on synapomorphies, the basic rationale
being that they were small Triassic theropods. Current research suggests Saltopus
is a more basal dinosauriform (e.g. Langer, 2004), and Avipes and Velocipes
to be avemetatarsalians of more dubious nature (Rauhut and Hungerbuhler, 2000).
Lukousaurus has not been recently restudied, but lacks several coelophysoid
apomorphies and is more likely crurotarsan. Elaphrosaurus was assigned
to Coelophysoidea (as Coelophysidae) by a couple authors (Paul, 1988; Novas,
1992) before it was realized to be a ceratosaur sensu lato (Holtz, 1994), which
has been the result of every published cladistic analysis. Paul (1988) also
believed spinosaurids were derived from coelophysids, but the description of
more complete spinosaurid specimens has led to their classification as tetanurines
(Sereno et al., 1996; Charig and Milner, 1997). Rauhut (2003) found Shuvosaurus
to fall out as a coelophysoid in his analysis, but the description of Effigia
(Nesbitt and Norell, 2006) led to its recognition as a crurotarsan.
Coelophysoidea defined- Sereno's (1998) and Padian et al.'s (1999) definitions
for Coelophysodea are basically equivalent as nearly every phylogeny agrees
Ceratosaurus and Carnotaurus are more closely related to each
other than either is to Coelophysis. The presence of Passer as
a specifier in Sereno's (in press) redefinition is useful, but Carnotaurus
seems superfluous, since there has never been a (Ceratosaurus (Passer
(Carnotaurus, Coelophysis))) topology suggested.
References- Huene, 1914. Beiträge zur geschichte der Archosaurier
[Contribution to the history of the archosaurs]. Geologie und Paläontologie
Abhandlungen. 13(7), 1-56
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Russell, 1984. A check list of the families and genera of North American dinosaurs.
Syllogeus. 53, 1-35.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Carroll, 1988. Vertebrate Paleontology and Evolution. W.H. Freeman and Company.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the
Early Jurassic Kayenta Formation of Arizona. J. Vert. Paleontol. 9, 125-136.
Novas, 1991. Relaciones filogeneticas de los dinosaurios teropodos ceratosaurios
[Phylogenetic relationships of ceratosaurian theropod dinosaurs]. 28(3-4), 401.
Novas, 1992. La evolucion de los dinosaurios carnivoros [The evolution of carnivorous
dinosaurs]. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico:
Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez",
Cuenca, Argentina. 126-163.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Charig and Milner, 1997. Baryonyx walkeri, a fish-eating dinosaur from
the Wealden of Surrey. Bulletin of the Natural History Museum of London (Geology).
53, 11-70.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. N. Jb. Geol. Paläont. Abh. 210(1),
41-83.
Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor,
Varricchio, Wilson and Wilson, 1998. A long-snouted predatory dinosaur from
Africa and the evolution of the spinosaurids. Science. 282(5392), 1298-1302.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature
of the major taxonomic categories of the carnivorous Dinosauria (Theropoda).
Journal of Vertebrate Paleontology, 19(1), 69-80.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey, 80 pages.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia.
15, 75-88.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contr. Mus. Palaeont. Univ. Mich. 31, 1-42.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska.
The Dinosauria Second Edition. University of California Press. 861 pp.
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria
Second Edition. University of California Press. 861 pp.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The
Dinosauria Second Edition. University of California Press. 861 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa
and its implications for the early evolution of theropods. Palaeontologia Africana.
41, 105-122.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article,
38 pp. DOI 10.1080/08912960600845791.
Nesbitt and Norell, 2006. Extreme convergence in the body plans of an early
suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of
the Royal Society B. 273, 1045-1048.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti
(Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications
for early theropod evolution. Zoological Journal of the Linnean Society. 151,
377-421.
unnamed possible coelophysoid (Galton, 1976)
Hettangian, Early Jurassic
Shuttle Meadow Formation, Connecticut, US
Material- tooth
Comments- Galton (1976) referred this to Coelophysis sp., but
Coelophysis teeth are not known to be diagnostic.
Reference- Galton, 1976. Prosauropod dinosaurs (Reptilia: Saurischia)
of North America. Postilla. 169:1-98.
undescribed possible coelophysoid (Ford, DML 1998)
Early Jurassic
Hanson Formation, Antarctica
Material- teeth
Comments- Although described as 'halticosaurid' by Ford, coelophysoid
teeth are not known to be diagnostic.
Reference- http://dml.cmnh.org/1998Aug/msg00810.html
unnamed coelophysoid (Cunningham, Hungerbuhler, Chatterjee and McQuilkin,
2002)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US
Material- (TTUP coll.) tibia
Comments- Cunningham et al. (2002) identified this specimen as an ornithischian,
but Nesbitt et al. (2007) noted it has theropod synapomorphies like a fibular
crest, and shares a subrectangular distal outline with a small posterolateral
process with coelophysoids. It is indistinguishable from Coelophysis
and Gojirasaurus.
References- Cunningham, Hungerbuhler, Chatterjee and McQuilkin, 2002.
Late Triassic vertebrates from the Patricia site near Post, Texas. Journal of
Vertebrate Paleontology. 22(supplement to 3). 47A.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
unnamed possible coelophysoid (Maisch and Matzke, 2003)
Bathonian-Callovian, Middle Jurassic
Toutunhe Formation, Xinjiang, China
Material- (SGP 2000/2) (~3 m) distal fibula
(SGP 2001/5) (~3 m) tooth
Comments- Maisch et al. (2001) labeled the fibula as a coelurosaur. Maisch
and Matzke (2003) note the fibula is most similar to Sarcosaurus and
Megapnosaurus, based on the anteroproximal inclination of the distal
articular facet. The tooth was assigned to Coelophysoidea based on the large
number of serrations (33-37/mm), which is comparable to Coelophysis and
Liliensternus.
Reference- Maisch and Matzke, 2003. Theropods (Dinosauria, Saurischia)
from the Middle Jurassic Toutunhe Formation of the southern Junggar Basin, NW
China. Palaeontologische Zeitschrift. 77(2):281-292.
Sarcosaurus Andrews, 1921
?= "Liassaurus" Welles, Powell and Pickering vide Pickering, 1995
S. woodi Andrews, 1921
= Magnosaurus woodi (Andrews, 1921) Huene, 1932
?= "Liassaurus huenei" Welles, Powell and Pickering vide Pickering,
1995
Sinemurian, Early Jurassic
Lower Lias, England
Holotype- (BMNH R4840/1) (robust) (2.89 m, 70 kg) partial posterior dorsal
vertebra, incomplete ilium (248 mm), proximal pubis, ischial fragment, femur
(315 mm)
Referred- ?(Warwick Museum coll.) two dorsal centra, ilial fragment,
partial pubis, femora, tibia, partial tibia, distal fibula, distal metatarsal
II, distal metatarsal III, distal metatarsal IV, proximal pedal phalanx (Huene,
1932)
Diagnosis- (from Welles, 1984) ilium differs from Dilophosaurus wetherilli
in dorsal edge less convex over pubic peduncle; lower preacetabular process;
smaller supracetabular crest; narrower ischial peduncle; femur differs from
Liliensternus liliensterni in less ventrally directed femoral head; more
proximally placed anterior trochanter; lower and longer fourth trochanter; less
sigmoid femoral shaft anteriorly.
Comments- Carrano and Sampson (2004) noted the ilium is similar to Dilophosaurus
while the femur resembles Liliensternus. Thus, though there may be no
strict apomorphies to diagnose Sarcosaurus, the combination of character
states is diagnostic and it is not a nomen dubium (contra Carrano and
Sampson).
Huene (1932) assigned the species to Magnosaurus, though this is certainly
incorrect as the latter is a megalosauroid.
"Liassaurus"- Pickering (1995) listed the nomen nudum
"Liassaurus huenei", which subsequent internet communication (2005)
has revealed to be the Warwick Museum specimen Huene (1932) referred to S.
woodi. Carrano and Sampson note it is extremely similar to S. woodi,
of similar size and contemporaneous. None of the characters listed in Pickering's
online diagnosis of "Liassaurus" are absent in S. woodi except
for the supposed absence of the trochanteric shelf, which is contradicted by
Carrano and Sampson, and (sexually?) dimorphic in coelophysoids in any case.
Thus, "Liassaurus" is here synonymized with Sarcosaurus woodi,
pending publication of the former by Pickering.
Relationships- Andrews (1921) originally assigned this genus to the Megalosauridae,
in which he included not only megalosauroids, but also Ceratosaurus and
carnosaurs. Specifically, Andrews felt the ilium was similar to Ceratosaurus
and Megalosaurus, while the anterior trochanter was more primitive than
Allosaurus, Megalosaurus and Ceratosaurus, though closest
to the latter. An assignment to such a broad Megalosauridae was standard throughout
the 1900's. Kurzanov (1989) referred it to his more restricted concept of Megalosauridae
(containing Megalosaurus and sinraptorids) without comment. Huene (1956)
referred it to his incorrectly formed family Coelurosauridae. Gauthier (1986)
recognized Sarcosaurus as a ceratosaur sensu lato on the basis of its
trochanteric shelf. This was elaborated on by Rowe (1989) and Rowe and Gauthier
(1990), who placed Sarcosaurus in Ceratosauria but outside their unnamed
equivalent of Coelophysidae based on several characters. Paul (1988) referred
Sarcosaurus to the Ceratosauridae. Welles (1984) was the first to suggest
coelophysoid relations, placing it in Coelophysidae with Coelophysis
(but not other currently recognized coelophysoids) without comment. Carrano
and Sampson (2004) described characters shared with coelophysoids, such as long
dorsal centra, an anteriorly facing pubic peduncle, small, spike-like anterior
trochanter, and notched, circular distal tibia. They also noted resemblence
to Liliensternus and Dilophosaurus, and referred Sarcosaurus
to Coelophysoidea. Although Carrano et al. (2005) could not recover any resolution
within Coelophysoidea in their cladistic analysis, when several characters are
changed to ordered, the results differ. If a standard coelophysoid topology
of (Dilophosaurus (Liliensternus (Coelophysis, Megapnosaurus)))
is enforced, Sarcosaurus is placed outside the Liliensternus +
Coelophysis + Megapnosaurus clade. Tykoski (2005) notes that Sarcosaurus
has a mix of coelophysoid and ceratosaur sensu stricto characters, and
found it to be the most basal coelophysoid in their analysis based on two characters-
supracetabular crest flares lateroventrally to hide dorsal acetabulum in lateral
view; anterior trochanter conical.
References- Andrews 1921. On some remains of a theropodous dinosaur from
the Lower Lias of Barrow-on-Soar. Annual Magazine of Natural History 9(8) 570-576.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monog. Geol. Pal. 4 (1) pts. 1 and 2, viii + 361 pp.
Huene, 1956. Paläontologie und Phylogenie der Niederen Tetrapoden [Paleontology
and Phylogeny of the Lower Tetrapods]. VEB Gustav Fischer Verlang, Jena. 1-716.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie 185 p. 85-180.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Kurzanov, 1989. O proiskhozhdenii i evolyutsii infraotryada dinozavrov Carnosauria
[Concerning the origin and evolution of the dinosaur infraorder Carnosauria].
Paleontologicheskiy Zhurnal. 1989(4), 3-14.
Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the
Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology.
9, 125-136.
Rowe and Gauthier, 1990. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.).
The Dinosauria. Univ. Cal. press. Berkeley.
Pickering, 1995. "Jurassic Park: Unauthorized Jewish Fractals in Philopatry,"
A Fractal Scaling in Dinosaurology Project, 2nd revised printing, Capitola,
California: 478 pp. [January 27, 1995].
Carrano and Sampson, 2004. A review of coelophysoids (Dinosauria: Theropoda)
from the Early Jurassic of Europe, with comments on the late history of the
Coelophysoidea. N. Jb. Geol. Palaont. Mh. 2004 (9): 537-558.
http://groups.yahoo.com/group/paleo_bio_dinosaur_ontology/message/8461
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), 835849.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Gojirasaurus Carpenter, 1997
= "Revueltoraptor" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
G. quayi Carpenter, 1997
= "Revueltoraptor lucasi" Hunt 1994 vide Nesbitt, Irmis and Parker,
2007
Norian, Late Triassic
Bull Canyon (=Cooper Canyon) Formation of the Dockum Group, New Mexico, US
Holotype- (UCM 47221; in part) (5.5 m) pubis (497 mm), tibia (469 mm)
Diagnosis- Provisionally indeterminate relative to Coelophysis bauri.
Comments- This specimen was originally briefly described and illustrated
as Procompsognathidae gen. et sp. indet. by Parrish and Carpenter (1986).
It was then described and named "Revueltoraptor lucasi" by Hunt (1994)
in his unpublished thesis, and called herrerasaurid A by Hunt et al. (1998).
It was finally officially named and described as the coelophysoid Gojirasaurus
quayi by Carpenter (1997). However, Nesbitt et al. (2005) find the holotype
to be chimaeric, which was elaborated on in their 2007 publication. They found
the dorsal vertebrae were referrable to Shuvosaurus and the pubis and
tibia referrable to a coelophysoid (contra the 2005 abstract where the dinosaurian
elements were relegated to Saurischia indet.). They could not assign
the other material (teeth, dorsal ribs, gastralia, chevron) or the several referred
specimens noted by Hunt (1994) (NMMNH P4666, P16607, P16656, P16946, P17134,
P17154, P17258, UMMP 7274) to a particular taxon of archosaur. The reassignment
of the dorsal material makes the sole apomorphy identified by Rauhut invalid
(mid/posterior dorsal vertebrae with taller neural spines than other coelophysoids),
and Gojirasaurus is presently indeterminate relative to Coelophysis
except for the greater robustness of the tibia (Nesbitt et al., 2007). This
may be size-related.
Although the name "Revueltoraptor lucasi" was originally used in thesis,
and thus not available for use in this website, it was later published by Nesbitt
et al. (2007). The name was first noted publically on the Dinosaur Mailing List
in 2000 by Gay, who noted several elements at the NMMNH were labeled as "Revueltoraptor".
These were specimens referred to the taxon by Hunt (1994), and are not identifiable
as Gojirasaurus.
Rauhut (2003) suggested Gojirasaurus may be synonymous with Shuvosaurus,
as a large Shuvosaurus premaxilla was found in the same deposits (identified
as Reptilia indet. by Parrish and Carpenter, 1986), and the tooth associated
with the Gojirasaurus holotype may not belong to it. However, Nesbitt
and Norell (2006) have recently demonstrated that Shuvosaurus is a crurotarsan
whose pubis and tibia differ markedly from Gojirasaurus'.
Relationships- Hunt (1994) and Hunt et al. (1998) referred this specimen
to Herrerasauridae based on the short dorsal centra, strap-like scapula and
elongate pubis. However, the dorsals are now assigned to Shuvosaurus,
the scapula is Archosauria indet., and all theropods have elongate pubes.
Parrish and Carpenter (1986) believed the taxon to be most closely related to
Liliensternus within their Procompsognathidae, in which they included
all coelophysoids. Carpenter (1997) concurred regarding the similarity to Liliensternus,
referring it to Coelophysoidea and possibly Coelophysidae. Rauhut's analysis
(2003) found Gojirasaurus to be a coelophysoid more closely related to
coelophysids than to Liliensternus. Yates' (2005) updated version of
Rauhut's analysis found Gojirasaurus to be in a polytomy with Megapnosaurus,
"M." kayentakatae and Segisaurus, more derived than
Coelophysis. Tykoski and Rowe (2004) could only say that Gojirasaurus
was a coelophysoid more derived than Dilophosaurus, which was also the
result of Carrano et al.'s (2005) analysis. Tykoski (2005) could find even less
resolution, with Gojirasaurus having an uncertain position within Coelophysoidea,
though in a polytomy with Zupaysaurus, Liliensternus and Coelophysis
in the majority rule tree. Finally, Ezcurra and Novas (2006) found it to be
the sister taxon of "Megapnosaurus" kayentakatae, outside Coelophysinae.
However, all of these analyses included Shuvosaurus and Archosauria indet.
remains in their Gojirasaurus OTUs, as they were published prior to Nesbitt
et al.'s reanalysis of the holotype. Thus any particular placement within Coelophysoidea
is questionable, and must be reexamined to determine if it was based on pubic
or tibial characters.
References- Parrish and Carpenter, 1986. A new vertebrate fauna from
the Dockum Formation (Late Triassic) of eastern New Mexico. In: Padian, K. (ed.).
The Beginning of the Age of Dinosaurs. Cambridge Univ. Press, NewYork. Pp.151-160.
Hunt, 1994. Vertebrate paleontology and biostratigraphy of the Bull Canyon Formation
(Chinle Group: Norian), east-central New Mexico with revisions of the families
Metoposauridae (Amphibia: Temnospondyli) and Parasuchidae (Reptilia: Archosauria).
Unpublished PhD Dissertation. Albuquerque, Univerrsity of New Mexico. 403 pp.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the
Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Carpenter, 1997. A giant coelophysoid (Ceratosauria) theropod from the Upper
Triassic of New Mexico, USA. Neues Jahrbuch fuer Geologie und Palaeontologie,
Abhandlungen. 205(2), 189-208.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from
the Western United States. Geobios 31, 4: 511-531.
http://dml.cmnh.org/2000Oct/msg00495.html
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The
Dinosauria Second Edition. University of California Press. 861 pp.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli,
a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate
Paleontology. 25(4), 835849.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur
record, part 3: Saurischians of North America. JVP 25(3) 96A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa
and its implications for the early evolution of theropods. Palaeontologia Africana.
41, 105-122.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article,
38 pp. DOI 10.1080/08912960600845791.
Nesbitt and Norell, 2006. Extreme convergence in the body plans of an early
suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of
the Royal Society B. 273, 1045-1048.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
Halticosauridae Bock, 1952
Halticosaurinae Bock, 1952 sensu Paul, 1988
Comments- Bock (1952) originally named this as a family of coelurosaurs
(sensu Huene) containing Liliensternus (as Halticosaurus liliensterni).
It is often ascribed to Huene (1956). Welles (1984) included Dilophosaurus,
Halticosaurus, Liliensternus and Longosaurus, while Chatterjee
(1993) and Hu (1993) also included the first two genera. Paul (1988) used a
subfamily Halticosaurinae to include Liliensternus and Dilophosaurus.
The family has never been defined with explicit synapomorphies, instead being
a receptacle for the larger coelophysoids which current analyses indicate form
a grade basal to coelophysids. Although Halticosaurus has never been
entered in a data matrix, Liliensternus and Dilophosaurus have
never formed a clade exclusive of Coelophysis in any published study.
Thus Halticosauridae has remained unused by most current workers. If Halticosaurus
is shown in the future to share synapomorphies with Liliensternus, Dilophosaurus,
Sarcosaurus or another taxon outside Coelophysidae, Halticosauridae should
be applied to the resulting clade.
References- Bock, 1952. Triassic reptilian tracks and trends of locomotive
evolution. Journal of Paleontology. 26(3), 395-433.
Huene, 1956. Paläontologie und Phylogenie der Niederen Tetrapoden [Paleontology
and Phylogeny of the Lower Tetrapods]. VEB Gustav Fischer Verlang, Jena. 1-716.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Chatterjee, 1993. Shuvosaurus, a new theropod: an unusual theropod dinosaur
from the Triassic of Texas. National Geographic Research and Exploration. 9(3),
274-285.
Hu, 1993. A new Theropoda (Dilophosaurus sinensis sp. nov.) from Yunnan,
China. Vertebrata PalAsiatica. 31(1), 65-69.
Halticosaurus Huene, 1907
H. longotarsus Huene, 1907
Middle Norian, Late Triassic
Middle Stubensandstein, Germany
Holotype- (SMNS 12353) dentary fragment (lost), teeth (lost), sixth (?)
cervical vertebra (57 mm), two cervical vertebrae (lost), anterior dorsal vertebra
(33 mm) (lost), dorsal centrum (43 mm), second sacral vertebra (35 mm) (lost),
third sacral vertebra (33 mm) (lost), proximal caudal vertebra (36 mm), proximal
humerus (lost), ilial fragments (lost), proximal femora (300+ mm), metatarsal
II (134 mm)
Referred- (HMN coll.) fragmentary remains (Huene, 1921)
Diagnosis- (after Welles, 1984) differs from Liliensternus liliensterni
in- shallower dentary; dentary more pointed anteriorly; shorter and taller mid
cervical vertebrae; sharp ventral keel on mid cervical vertebrae; narrower sacral
centra; lower third sacral vertebra; more distally placed anterior trochanter;
smaller distal condyles on metatarsal II which extend less far proximally on
shaft.
Comments- Norman (1990) and Rauhut and Hungerbuhler (2000) believe Halticosaurus
is indeterminate. However, Welles (1984) finds differences between comparable
taxa such as Liliensternus and Dilophosaurus.
Relationships- Huene (1908) originally assigned Halticosaurus
only to Dinosauria, then later (1909) to Saurischia. Romer's (1956) assignment
to Hallopodidae is incorrect, as Hallopus is a crurotarsan differing
from Halticosaurus and other theropods in having a proximally unprojected
anterior trochanter among other characters. Norman (1990) believed the remains
could be from a basal sauropodomorph or a theropod, though the sauropodomorph
characters seems to be untrue (cervical pleurocoels absent; two sacral vertebrae;
similar femur), and the theropod character primitive (elongate metatarsal).
Assignments to Compsognathidae (Zittel, 1911) and Coeluridae (Huene, 1920) are
similarly incorrect, as the low spike-like anterior trochanter is more primitive
than coelurosaurs. Most workers have assigned Halticosaurus to Podokesauridae
(Huene, 1914), an equivalent Procompsognathidae (Romer, 1966), or its eponymous
family Halticosauridae (Bock, 1952). These categories are equivalent to the
modern concepts of Coelophysoidea, with halticosaurids now thought to be a basal
grade of that clade. Between 1934 and 1984, such statements were usually based
on Liliensternus, then believed to be a species of Halticosaurus.
Rauhut and Hungerbuhler (2000) give the only modern account of H. longotarsus,
noting the material is very poorly preserved and most is not identifiable as
theropod. As the holotype was found with Sellosaurus gracilis remains,
some may be prosauropod. The proximal femora show a spike-like lesser trochanter
and downturned head, as in coelophysoids. They therefore think some of the type
may be coelophysoid. If this is true, the short cervical centrum excludes it
from Coelophysidae. However, Halticosaurus has never been included in a phylogenetic
analysis or described in detail since its discovery
References- Huene, 1908. Die Dinosaurier der Europäischen Triasformation
mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs
of the European Triassic formations with consideration of occurrences outside
Europe]. Geologische und Palaeontologische Abhandlungen Suppl. 1(1), 1-419.
Huene, 1909. Skizze zu einer Systematik und Stammesgeschichte der Dinosaurier
[Sketch of the systematics and origins of the dinosaurs]. Centralblatt für
Mineralogie, Geologie und Paläontologie. 1909, 12-22.
Zittel, 1911. Grundzüge der Paläontologie (Paläozoologie). II.
Abteilung. Vertebrata [Fundamentals of Paleontology (Paleozoology). Section
II. Vertebrata]. Druck und Verlag von R. Oldenbourg, München. 1-598.
Huene, 1914. Das natürliche System der Saurischia [The systematics of the
Saurischia]. Centralblatt für Mineralogie, Geologie und Paläontologie.
1914, 154-158.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien
[Phylogenetic results of some investigations of Triassic reptiles]. Zeitschrift
für Induktive Abstammungsund Vererbungslehre. 24, 159-163.
Huene, 1921. Coelurosaurier-Reste aus dem obersten Keuper von Halberstadt. Cbl.
Min. Geol. Palaeontol. 1921(10), 315-320.
Bock, 1952. Triassic reptilian tracks and trends of locomotive evolution. Journal
of Paleontology. 26(3), 395-433.
Huene, 1956. Paläontologie und Phylogenie der Niederen Tetrapoden [Paleontology
and Phylogeny of the Lower Tetrapods]. VEB Gustav Fischer Verlang, Jena. 1-716.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press,
Chicago. 1-468.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie 185 p. 85-180.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. Univ. Cal. press. Berkeley.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia.
15, 75-88.
Sarcosaurus? andrewsi Huene,
1932
= Magnosaurus woodwardi Huene, 1932
= Megalosaurus andrewsi (Huene, 1932) Waldman, 1974
Sinemurian, Early Jurassic
Lower Lias, England
Holotype- (BMNH R3542) tibia (445 mm)
Diagnosis- Provisionally indeterminate relative to Sarcosaurus woodi.
Comments- Carrano and Sampson (2004) found it to be indistinguishable
from the referred specimen of Sarcosaurus woodi except for size, and
suggested the two species might be synonyms. The specimen was accidentally made
the type of two species simultaneously by Huene (1932), who later (1956) used
S. andrewsi for it.
References- Woodward, 1908. Note on a megalosaurian tibia from the Lower
Lias of Wilmcote, Warwickshire. Annals and Magazine of Natural History. ser.
8 (1) 257-259.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monog. Geol. Pal. 4 (1) pts. 1 and 2, viii + 361 pp.
Huene, 1956. Paldontologie und Phylogenie der niederen Tetrapoden. Fischer-Verlag,
Jena.
Carrano and Sampson, 2004. A review of coelophysoids (Dinosauria: Theropoda)
from the Early Jurassic of Europe, with comments on the late history of the
Coelophysoidea. N. Jb. Geol. Palaont. Mh. 2004 (9): 537-558.
unnamed clade (Dilophosaurus wetherilli + Coelophysis bauri)
Diagnosis- (after Tykoski, 2005) skull elongate (>3 times longer than
tall); antorbital fenestra elongate (>25% skull length); premaxillary tooth
row ends anterior to external naris; elongate posteromedial process on premaxilla;
premaxilla and maxilla loosely articulated; premaxillary and maxillary alveolar
margins do not contact; anterior tip of maxillary alveolar margin curves mediodorsally;
first maxillary alveolus opens anteroventrally; anteromedial maxillary process
elongate; dorsal edge of anterior dentary raised; most premaxillary teeth lack
serrations; anterior dentary teeth enlarged; narrow cervical epipophyses which
project posterolaterally; no axial diapophyses; axial parapophyses reduced or
absent; pubic peduncle of ilium with two facets- facing anteriorly and ventrally;
distal tips of pubes lack contact; nasal process of premaxilla comprises >50%
of external naris' dorsal edge; frontals elongated and triangular; lateral lamina
of lacrimal sinuous and extends anteriorly past medial lamina; anterior pleurocoels
in cervical vertebrae; elongate anterior dorsal centra (>1.33 height of anterior
articular surface); elongate posterior dorsal centra (>1.33 height of anterior
articular surface); ventral surface of caudal centra with narrow, sharp longitudinal
groove; posterior scapular margin nearly straight over majority of length; distal
carpals I and II fused; phalanx I-1 elongated (> 100% metacarpal I length);
manual digit II longer than digit III; ischial antitrochanter large, protrudes
anteriorly into acetabulum; obturator notch in ischium; oblique ridge on proximomedial
fibula that overlaps groove.
Dilophosauridae Charig and Milner, 1990
= Dilophosaurinae Charig and Milner, 1990 sensu Madsen and Welles, 2000
Diagnosis- (after Smith et al., 2007) contribution of the premaxillary
posterodorsal process to a blade-like nasal crest; extension of the antorbital
fossa onto the
lateroventral side of the nasal; presence of a nasolacrimal crest.
Comments- Although historically various taxa (Liliensternus, Halticosaurus,
etc.) have been proposed to be more closely related to Dilophosaurus
than to Coelophysis or tetanurines (e.g. Welles, 1984; Paul, 1988), these
were not supported by phylogenetic analyses. The first analysis to recover such
sister taxa to Dilophosaurus was that of Yates (2005), which found a
clade containing Dilophosaurus, Zupaysaurus and Dracovenator
to be sister to Ceratosauria+Tetanurae. Smith et al. (2007) later found a clade
containing "Dilophosaurus" sinensis, Dracovenator, Cryolophosaurus
and Dilophosaurus (but not Zupaysaurus, which was slightly more
basal). Zupaysaurus has been assigned to Coelophysidae in other recent
analyses (Carrano et al., 2005; Tykoski, 2005; Ezcurra and Novas, 2006). Smith
et al.'s dilophosaur clade was also sister to Ceratosauria+Tetanurae. Both Yates'
and Smith et al.'s analyses could place their dilophosaur clades in Coelophysoidea
with few added steps (1 in Yates; 6 in Smith et al.).
Dilophosauridae was first used by Charig and Milner (1990) to refer to Paul's
(1988) informal dilophosaurs, which he actually used the subfamiliy Halticosaurinae
for. 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. While neither Yates nor Smith et al. used a name for their dilophosaur
clade, Dilophosauridae seems appropriate and is used here.
References- Paul, 1988. Predatory Dinosaurs of the World. Simon &
Schuster, New York.
Charig and Milner, 1990. The systematic position of Baryonyx walkeri,
in the light of Gauthier's reclassification of the Theropoda. in Carpenter and
Currie (eds.). Dinosaur Systematics: Approaches and Perspectives. Cambridge
University Press, Cambridge. 127-140.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey, 80 pages.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli,
a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate
Paleontology. 25(4), 835849.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa
and its implications for the early evolution of theropods. Palaeontologia Africana.
41, 105-122.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article,
38 pp. DOI 10.1080/08912960600845791.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti
(Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications
for early theropod evolution. Zoological Journal of the Linnean Society. 151,
377-421.
"Dilophosaurus" sinensis
Hu, 1993
= Dilophosaurus sinensis Dong, Hisa and Azuma, 1990
Hettangian-Pliensbachian, Early Jurassic
Lower Lufeng Formation, Yunnan, China
Holotype- (Kunming Municipal Museum KMV 8701) (5.5 m) skull (525 mm),
lower jaw (487 mm), nine cervical vertebrae, cervical ribs, fifteen dorsal vertebrae,
four sacral vertebrae, thirty-six caudal vertebrae, chevrons, scapulae, coracoids,
distal clavicle, humerus, radius, ulna, metacarpals, ilium, pubis, ischium,
femur (587 mm), tibia, fibula, astragalus, calcaneum, distal tarsal IV, metatarsus,
pes
Comments- Lamanna et al. (1999) doubt this species is referrable to Dilophosaurus
based on differences in premaxillary shape, tooth count, lack of antorbital
tooth row, shape of infratemporal fenestra and squamosal and size and position
of external mandibular fenestra. Dong (2003) believes this taxon is similar
to, and perhaps synonymous with Sinosaurus triassicus, though without
citing supporting evidence. Smith et al. (2007) included it in their phylogenetic
analysis, where it emerged as the most basal dilophosaurid. Unless Dracovenator
and Cryolophosaurus are synonymized with Dilophosaurus as well,
"D." sinensis deserves a new genus name.
References- Hu, 1993. A new Theropoda (Dilophosaurus sinensis
sp. nov.) from Yunnan, China. Vertebrata PalAsiatica. 31(1), 65-69.
Lamanna, Smith, You, Holtz and Dodson, 1999. A reassessment of the Chinese theropod
dinosaur Dilophosaurus sinensis. JVP 18(3) 57A.
Dong, 2003. Contributions of new dinosaur materials from China to dinosaurology.
Memoir of the Fukui Prefectural Dinosaur Museum. Number 2 pp. 123-131.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti
(Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications
for early theropod evolution. Zoological Journal of the Linnean Society. 151,
377-421.
Sinosaurus Young, 1948
S. triassicus Young, 1948
Sinemurian, Early Jurassic
Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Holotype- (IVP AS V34) two maxillary fragment, four maxillary teeth,
jaw fragment, three teeth
Paratypes- (IVP AS V35, V36, V37, V48) teeth
Hettangian, Early Jurassic
Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
Paratype- (IVP AS V38) teeth
Hettangian-Sinemurian, Early Jurassic
Lufeng Formation, Yunnan, China
Referred- (CUP 2001) tooth (Simmons, 1965)
(CUP 2002) tooth (Simmons, 1965)
(CUP 2003) dorsal centrum (Simmons, 1965)
(CUP 2004) tooth (Simmons, 1965)
(CUP 2005) tooth (Simmons, 1965)
(CUP 2095) dorsal centrum (Simmons, 1965)
(CUP 2096) three tooth fragments (Simmons, 1965)
(CUP 2097) anterior jaw fragment, teeth (Simmons, 1965)
(CUP 2098) cervical centrum, two dorsal centra (Simmons, 1965)
Comments- Dong (2003) believes this taxon is similar to, and perhaps
synonymous with "Dilophosaurus" sinensis, though without citing
supporting evidence. The postcrania referred to the taxon by Young (1948; 1951)
are sauropodomorph (Walker, 1964), later assigned to Jingshanosaurus
(Zhang and Yang, 1995), but the latter referral was doubted by Galton and Upchurch
(2004).
References- Young, 1948. On two new saurischians from Lufeng, Yunnan.
Bull. Geol. Soc. China 28 75-90, 6 figs., 8 pls.
Young, 1951. The Lufeng saurischian fauna in China. Palaeontologica Sinica.
C (13), 1-96.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and
the origin of carnosaurs. Philos. Trans. R. Soc. London B 248, 53-134.
Simmons, 1965. The non-therapsid reptiles of the Lufeng Basin, Yunnan, China.
Fieldiana Geol. 15, 1-93.
Zhang and Yang, 1995. A complete osteology of Prosauropoda in Lufeng Basin Yunnan
China. Jingshanosaurus. Yunnan Science and Technology Publishing House,
Kunming. 100 pp.
Dong, 2003. Contributions of new dinosaur materials from China to dinosaurology.
Memoir of the Fukui Prefectural Dinosaur Museum. Number 2 pp. 123-131.
Galton and Upchurch, 2004. Prosauropoda. In Weishampel, Dodson and Osmolska.
The Dinosauria Second Edition. University of California Press. 861 pp.
unnamed clade (Dracovenator regenti + Cryolophosaurus ellioti + Dilophosaurus
wetherilli)
Diagnosis- (after Smith et al., 2005) low height/length ratio of the
premaxilla below the external naris; slot-shaped foramen on the lateral face
of the premaxilla at the base of the nasal process (unknown in "Dilophosaurus"
sinensis); presence of erect, tab-like dorsal processes on the articular
(unknown in "Dilophosaurus" sinensis).
Dracovenator Yates, 2005
D. regenti Yates, 2005
Hettangian-Sinemurian, Early Jurassic
Upper Elliot Formation
Holotype- (BP/1/5243) premaxillae, maxillary fragment, two dentary fragments,
partial surangular, partial angular, partial prearticular, articular, teeth
Paratype- ?(BP/1/5278) (juvenile) premaxillae, incomplete maxillae, partial
nasals, partial dentaries, teeth (Munyikwa and Raath, 1999)
Diagnosis- (after Yates, 2005) a large bilobed fossa surrounding a large
lateral premaxillary foramen that is connected to the alveolar margin by a deep
narrow channel; a deep, oblique notch on the lateral surface of the articular,
separating the retroarticular process from the posterior margin of the glenoid;
particularly well-developed dorsal, tab-like processes on the articular, one
on the medial side, just posterior to the opening of the chorda tympanic foramen
and the other on the lateral side on the anterolateral margin of the fossa for
the m. depressor mandibulae.
References- Munyikwa and Raath, 1999. Further material of the ceratosaurian
dinosaur Syntarsus from the Elliot Formation (Early Jurassic) of South
Africa. Palaeontologia Africana. 35:55-59.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa
and its implications for the early evolution of theropods. Palaeontologia Africana.
41: 105122.
unnamed clade (Cryolophosaurus ellioti + Dilophosaurus wetherilli)
Diagnosis- (after Smith et al., 2005) lateral groove along the posterior
end of the surangular.
Cryolophosaurus Hammer and
Hickerson, 1994
= "Elvisaurus" Holmes, 1993
C. ellioti Hammer and Hickerson, 1994
Rhaetian-Toarcian, Late Triassic-Early Jurassic
Hanson Formation, Antarctica
Holotype- (FMNH PR1821) (~6.5 m; 465 kg) (partial skull ~460 mm) two
maxillary fragments, nine maxillary teeth, posterior skull, posterior mandibles,
sixth cervical central fragment, seventh cervical vertebra (118 mm), eighth
cervical vertebra (108 mm), ninth cervical vertebra, tenth cervical vertebra,
several posterior cervical ribs, several anterior dorsal vertebrae, most mid
and posterior dorsal vertebrae (114, 125, 115, 117 mm), several dorsal ribs,
fifth sacral vertebra (89 mm), partial first caudal centrum, two mid caudal
vertebrae (100 mm), three mid caudal centra, many partial and complete caudal
vertebrae, three chevrons, partial humerus, proximal radius, proximal ulna,
partial ilium, proximal pubis, ischia (one distal), incomplete femora (769 mm),
proximal tibia, distal tibia, distal fibula, astragalus, calcaneum
Diagnosis- (after Smith et al., 2007) large, anterodorsally curving midline
crest with fluted rostral and caudal surfaces formed by dorsal expansions of
the lacrimals; complete constriction across the infratemporal fenestra formed
by the squamosal and jugal; extremely elongate cranial processes on the cervical
ribs.
Comments- Smith et al. (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.
References- Holmes, 1993. Prehistoric Times.
Hammer and Hickerson, 1994. A crested theropod dinosaur from Antarctica. Science.
264:828-830.
Smith, Hammer and Currie, 2005. Osteology and phylogenetic relationships of
Cryolophosaurus ellioti (Dinosauria: Theropoda): Implications for basal
theropod evolution. JVP. 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.
Dilophosaurus Welles, 1970
D. wetherilli (Welles, 1954) Welles, 1970
= Megalosaurus wetherilli Welles, 1954
= Dilophosaurus breedorum Pickering, 1995
= Dilophosaurus breedorum Welles, 1995 vide Welles and Pickering, 1999
Hettangian, Early Jurassic
Silty Facies Member of Kayenta Formation, Arizona, US
Holotype- (UCMP 37302) (young subadult) (6.03 m, 283 kg) skull (523 mm),
atlas (19 mm), axial intercentrum (17 mm), axis (56 mm), third cervical vertebra
(~75 mm), fourth cervical vertebra (80 mm), fifth cervical neural arch, sixth
cervical vertebra (86 mm), seventh cervical neural arch, eighth cervical neural
arch (~87 mm), posterior ninth cervical centrum (~88 mm), posterior tenth cervical
centrum (~80 mm), first dorsal centrum (73 mm), second dorsal vertebra (67 mm),
third dorsal vertebra (~62 mm), fourth dorsal vertebra (67 mm), fifth dorsal
vertebra (74 mm), sixth dorsal vertebra (70 mm), seventh dorsal vertebra (78
mm), eighth dorsal vertebra (84 mm), ninth dorsal vertebra (82 mm), tenth dorsal
vertebra (~88 mm), eleventh dorsal vertebra (86 mm), twelfth dorsal vertebra
(90 mm), thirteenth dorsal vertebra (70 mm), dorsal ribs, partial first sacral
vertebra, second sacral vertebra (69 mm), third sacral vertebra (67 mm), fourth
sacral vertebra (67 mm), fifth sacral vertebra (70 mm), first caudal vertebra
(69 mm), second caudal vertebra (65 mm), third caudal vertebra (65 mm), caudals
4-44, chevrons 2-36, scapulae (375 mm), coracoids (105 mm high, 185 mm long),
humeri (285, 270 mm), radii (180, 192 mm), ulnae (205, 209 mm), radiale, intermedium,
ulnare, distal carpals I-III, 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), ilia (370 mm), pubes (485 mm),
ischia (340 mm), femur (557 mm), tibia (555 mm), fibula (518 mm), astragalus
(92 mm wide, 77 mm tall), calcaneum, two distal tarsals, metatarsal I (95 mm),
phalanx I-I (68 mm), pedal ungual I (55 mm), metatarsal II (250 mm), phalanx
II-1 (103 mm), phalanx II-2 (79 mm), pedal ungual II (73 mm), metatarsal III
(300 mm), phalanx III-1 (110 mm), phalanx III-2 (84 mm), phalanx III-3 (70 mm),
pedal ungual III (70 mm), metatarsal IV (254 mm), phalanx IV-1 (72 mm), phalanx
IV-2 (57 mm), phalanx IV-3 (46 mm), phalanx IV-4 (38 mm), pedal ungual IV (45
mm), metatarsal V (126 mm)
Paratypes- (UCMP 3703) (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
(UCMP 77270; holotype of Dilophosaurus breedorum) (6.81 m; adult) skull
(590 mm)
Referred- (MNA P1.97) tooth (Gay, 2001)
(MNA P1.101) proximal pubis (Gay, 2001)
(MNA Pl.102) (juvenile) distal fibula (Gay, 2001)
(MNA P1.109) (juvenile) femur (281 mm) (Gay, 2001)
(MNA P1.111) dentary fragments (Gay, 2001)
(MNA P1.116) manual phalanx III-1 (48 mm) (Gay, 2001)
(MNA P1.122) ischial fragment (Gay, 2001)
(MNA P1.131) pedal phalanx III-1 (120 mm) (Gay, 2001)
(MNA P1.135) dorsal centrum (83 mm), partial neural spine (Gay, 2001)
(MNA P1.138) dorsal centrum (45 mm) (Gay, 2001)
(MNA P1.141) manual phalanx III-2 (60 mm) (Gay, 2001)
(MNA P1.142) partial rib (Gay, 2001)
(MNA P1.154) proximal pubis (Gay, 2001)
(MNA P1.160) proximal femur (~560 mm) (Gay, 2001)
(MNA P1.161) distal femur (Gay, 2001)
(MNA P1.176) dorsal centrum (31 mm) (Gay, 2001)
(MNA P1.177) dorsal centrum (85 mm), partial neural spine (Gay, 2001)
(MNA P1.247) partial rib (Gay, 2001)
(MNA P1.248) pubis (570 mm) (Gay, 2001)
(MNA P1.530) distal fibula (Gay, 2001)
(MNA P1.539) distal fibula (Gay, 2001)
(MNA P1.3181) (juvenile) tooth fragment, distal humerus, distal fibula (Gay,
2001)
(TMM 43646) (~5.1 m; young subadult) (Tykoski, 2005)
(TMM 43662-2) (gracile adult) proximal femur (Tykoski, 2005)
(UCMP 130053) incomplete skeleton (Clark and Fastovsky, 1986)
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.
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).
References- Welles, 1954. New Jurassic dinosaur from the Kayenta formation
of Arizona. Bull. Geol. Soc. Amer. 65 591-598, 1 pl., 2 tables.
Welles, 1983. Two centers of ossification in a theropod astragalus. Journal
of Paleontology 57 p. 401.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Clark and Fastovsky, 1986. Vertebrate biostratigraphy of the Glen Canyon Group
in northern Arizona. The Beginning of the Age of the Dinosaurs: Faunal change
across the Triassic-Jurassic boundary. in Fraser and Sues (eds.). Cambridge
University Press. 285-301.
Pickering, 1995. "Jurassic Park: Unauthorized Jewish Fractals in Philopatry,"
A Fractal Scaling in Dinosaurology Project, 2nd revised printing, Capitola,
California: 478 pp. [January 27, 1995].
Welles and Pickering, 1999. An Extract From: Archosauromorpha: Cladistics and
Osteologies. 70 pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Gay, 2001. Evidence for sexual dimorphism in the Early Jurassic theropod dinosaur,
Dilophosaurus and a comparison with other related forms. JVP 21(3) 53A.
Gay, 2001. New specimens of Dilophosaurus wetherilli (Dinosauria: Theropoda)
from the early Jurassic Kayenta Formation of northern Arizona. Western Association
of Vertebrate Paleontologists annual meeting volume. Mesa, Arizona.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
unnamed clade (Liliensternus liliensterni + Coelophysis bauri)
= Coelophysidae sensu Sereno, 1998
Definition- (Coelophysis bauri + Procompsognathus triassicus)
(modified)
Diagnosis- (after Tykoski, 2005) alveolar ridge on maxilla; deep ventral
fossa on ectopterygoid; low post-axial cervical neural spines; ischiopubic ratio
<66%; proximal end of metatarsal III with ventral boss; anterior ramus of
maxilla <10% maxillary length; more than 19 maxillary teeth; anterior margin
of antorbital fossa squared; anterior lacrimal process longer than ventral process;
cervical ribs fused to vertebrae in adults.
"Comanchesaurus" Hunt, 1994
vide Nesbitt, Irmis and Parker, 2007
"C. kuesi" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US
Material- (NMMNH P4569) dorsal centra, proximal femur, partial astragalus,
metatarsal fragments, phalanges
Comments- NMMNH P4569 was originally seen as a coelophysoid by Lucas
et al. (1985) and Hunt and Lucas (1989), though Murray and Long (1989) and Long
and Murray (1995) referred it to Sauropodomorpha. Hunt (1994) named this "Comanchesaurus
kuesi" in his unpublished thesis, assigning it to Herrerasauridae. It was
called herrerasaurid B by Hunt et al. (1998). Nesbitt et al. (2007) confirmed
a saurischian identity based on the dorsal astragalar basin, but noted the astragalar
morphology was closer to coelophysids than to Herrerasaurus or other
taxa. It may therefore be a coelophysoid, as originally believed.
Nesbitt et al. further noted the fragmentary remains referred to "Comanchesaurus"
are indeterminate.
Although the name "Comanchesaurus kuesi" was originally used in thesis,
and thus not available for use in this website, it was later published by Nesbitt
et al. (2007).
References- Lucas, Hunt and Bennett, 1985. Triassic vertebrates from
eastcentral New Mexico in the Yale Peabody Museum. New Mexico Geological Society
Guidebook. 36, 199203.
Hunt and Lucas, 1989. Late Triassic vertebrate localities in New Mexico. pp.
72-101. in Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American
Southwest. New Mexico Museum of Natural History, Albuquerque.
Murry and Long, 1989. Geology and paleontology of the Chinle Formation, Petrified
Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils
of the southwestern Upper Triassic. pp. 29-64. in Lucas & A. P. Hunt (eds).
Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of
Natural History, Albuquerque.
Hunt, 1994. Vertebrate paleontology and biostratigraphy of the Bull Canyon Formation
(Chinle Group: Norian), east-central New Mexico with revisions of the families
Metoposauridae (Amphibia: Temnospondyli) and Parasuchidae (Reptilia: Archosauria).
Unpublished PhD Dissertation. Albuquerque, Univerrsity of New Mexico. 403 pp.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the
Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from
the Western United States. Geobios 31, 4: 511-531.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
Dolichosuchus Huene, 1932
D. cristatus Huene, 1932
Early-Middle Norian, Late Triassic
Lower or Middle Stubsandstein, Germany
Holotype- (BMNH R38058) tibia (330 mm)
Diagnosis- Provisionally indeterminate relative to Liliensternus liliensterni.
Comments- Huene (1932) originally assigned this genus to Hallopodidae.
This has a large cnemial crest and fibular crest, showing it is theropod. Welles
(1984) found the differences from Liliensternus to be insignificant.
Rauhut and Hungerbuhler (2000) note close resemblence to Liliensternus
and Dilophosaurus, suggesting it is probably a coelophysoid.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monog. Geol. Pal. 4 (1) pts. 1 and 2, viii + 361
pp.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie 185 p. 85-180.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia
15, 75-88.
Procompsognathidae Nopcsa, 1923 sensu Huene, 1929
Comments- Huene (1929) separated Procompsognathus from podokesaurids
as a new family Procompsognathidae, which was not followed by many authors through
the 1900's. Huene (1932) included Pterospondylus as well, which was often
viewed as a synonym of Procompsognathus. Romer (1966) and others have
incorrectly used this as a senior synonym of Podokesauridae, including Avipes,
Coelophysis, Dolichosuchus, Halticosaurus, Lukousaurus,
Podokesaurus, Procompsognathus, Saltopus, Scleromochlus,
Spinosuchus, Trialestes and Velocipes in the family. Steel
(1970) only included Procompsognathus and Halticosaurus. Welles
(1984) and Paul (1988) both only included Procompsognathus in the family.
Procompsognathidae has not been used since the 1980's, as Procompsognathus
has most often been viewed as a coelophysoid incertae sedis or coelophysid,
and was even made an internal specifier of Coelophysidae by Sereno (1998). Yet
if the genus is a basal dinosauriform as Allen (2004) believes, the family may
be justified again.
References- Nopcsa, 1923. Die Familien der Reptilien [The families of
reptiles]. Forschritte der Geologie und Palaeontologie. 2, 1-210.
Huene, 1929. Kurze Übersicht über die Saurischia und ihre natürlichen
Zusammenhänge [A brief survey of the Saurischia and their natural context].
Paläontologische Zeitschrift. 11, 269-273.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monog. Geol. Pal. 1, pts 1 and 2, V111 + 361 pp.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press,
Chicago. 1-468.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia
of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 1-87.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und
Paläontologie, Abhandlungen. 210(1), 41-83.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP
24(3).
Procompsognathinae Nopcsa, 1923
Definition- (Procompsognathus triassicus <- Coelophysis
bauri) (Sereno, in press; modified from Sereno, 1998)
Comments- Nopcsa (1923) named this subfamily within Compsognathidae,
separate from Compsognathinae and Ornithomiminae. This was not followed, though
Sereno (1998) later used it as a subfamily of Coelophysidae to contain Procompsognathus
and Segisaurus, separate from the coelophysines Coelophysis and
Megapnosaurus. This was not found in future studies (e.g. Tykoski, 2005;
Ezcurra and Novas, 2006), where Procompsognathus instead has a more unstable
position more derived than Dilophosaurus. This makes the utility of Procompsognathinae
questionable and the application of Sereno's definition to any taxon except
Procompsognathus itself impossible. If Procompsognathus is non-dinosaurian
as Allen (2004) suggests, Procompsognathinae may have some use depending on
its exact relationships.
References- Nopcsa, 1923. Die Familien der Reptilien [The families of
reptiles]. Forschritte der Geologie und Palaeontologie. 2, 1-210.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und
Paläontologie, Abhandlungen. 210(1), 41-83.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP
24(3).
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article,
38 pp. DOI 10.1080/08912960600845791.
Procompsognathus Fraas,
1913
P. triassicus Fraas, 1913
= Hallopus "celerrimus" Fraas, 1912
Middle Norian, Late Triassic
Middle Stubensandstein, Germany
Holotype- (SMNS 12591) (.96 m) (robust) skull (associated?), partial
cervical vertebrae 5-10, partial cervical ribs, first dorsal vertebra, second
dorsal vertebra (~12.6 mm), third dorsal vertebra (~13.6 mm), fourth dorsal
vertebra (~13.6 mm), fifth dorsal vertebra, sixth dorsal vertebra (15.8 mm),
seventh dorsal vertebra (16.2 mm), eighth dorsal vertebra (16.2 mm), ninth dorsal
vertebra (16.8 mm), tenth dorsal vertebra (16.4 mm), partial dorsal ribs, thirteen
anterior caudal vertebrae 15 mm), scapulocoracoid (43+ mm), radius, ulna (34.2
mm), radiale?, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, partial
phalanx II-1, incomplete metacarpal III, partial metacarpal IV, partial ilium,
pubes (88+, 86+ mm), femora (92.5, 93.1 mm), tibia (112.6 mm), proximal tibia,
fibula, astragalus, metatarsal I (10.7 mm), phalanx I-1 (12.3 mm), pedal ungual
I (7.3 mm), metatarsal II (58 mm), phalanx II-1 (15.9 mm), phalanx II-2 (12.8
mm), pedal ungual II (10.6 mm), metatarsal III (69.4 mm), phalanx III-1 (17.1
mm), phalanx III-2 (15.2 mm), phalanx III-3 (12 mm), pedal ungual III (9.7 mm),
metatarsal IV (68.8 mm), phalanx IV-1 (~6.8 mm), phalanx IV-2 (8.7 mm), phalanx
IV-3 (7.2 mm), phalanx IV-4 (5.2 mm), pedal ungual IV (7.2 mm), metatarsal V
(23 mm) (Huene 1921, Ostrom 1981, Sereno and Wild 1992, Chatterjee 1993)
Diagnosis- (after Rauhut, 2000) scapula more slender than Coelophysis
bauri.
Comments- Hallopus "celerrimus" is probably a synonym
of Procompsognathus triassicus, according to Molnar (pers. comm.). He
suspects this based on the fact they come from the same locality and beds, were
proposed by the same person, and Fraas never used the name Hallopus "celerrimus"
after he published Procompsognathus triassicus.
Sereno and Wild (1992) considered the skull to belong to Saltoposuchus connectens,
while Chatterjee (1993) believed it to be theropod though the latter evidence
is not published yet.
A detailed description of the reprepared skull is in progress by Chatterjee
(pers. comm. to Rauhut, 1996), so Rauhut and Hungerbuhler (2000) do not consider
the skull in their commentary. They note that the slightly bowed pubis with
rectangular apron and absence of a pubic foot in this specimen are symplesiomorphies,
so do not help resolve relationships (contra Ostrom 1981, Sereno and Wild 1992).
They think the elongate dorsal vertebrae and triangular dorsal transverse processes
may indicate ceratosaurian or coelophysoid affinities. It is provisionally regarded
as valid based on an elongate hindlimb (tib/fem 1.2, mtIII/fem .74). I do not
find this character convincing, as Podokesaurus has similar ratios (tib/fem
1.21, mtIII/fem .76) and Coelophysis can also have similar proportions
(tib/fem 1.17, mtIII/fem .71). Still, I feel there is a good possibility future
work will provide valid synapomorphies.
Allen (2004) believes the skull belongs to the same taxon as the postcrania,
and based on cladistic analyses both including and excluding the former, places
Procompsognathus as a non-dinosaurian avemetatarsalian. However, the
evidence for this reinterpretation has not yet been published.
References- Fraas, 1912. Jahresh. Ver. Naturk. Wuerttemberg. 68:66-67.
Fraas, 1913. Die neuesten Dinosaurierfunde in der schwabischen Trias. Naturwissenschaften
45: 1097-1100.
Ostrom, 1981. Procompsognathus - theropod or thecodont?. Palaeontographica.
Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie,
Stratigraphie 175 p. 179-195.
Sereno and Wild, 1992. Procompsognathus: Theropod, ``thecodont'' or both?.
Journal of Vertebrate Paleontology 12 p. 435-458.
Chatterjee, 1993. Procompsognathus from the Triassic of Germany is not
a crocodylomorph. Abstract, Journal of Vertebrate Paleontology, 13(3): 29A.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia
15, 75-88.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP
24(3).
Protoaviformes Chatterjee, 1991
= Praeornithurae Kurochkin, 1995
= Protoaviornithes Kurochkin, 1995
Protoavidae Chatterjee, 1991
Protoavis Chatterjee, 1991
P. texensis Chatterjee, 1991
Norian, Late Triassic
Bull Canyon Formation of Dockum Group, Texas, US
Holotype- (TTU P 9200)
unassigned- premaxilla, lacrimal(?), squamosal, quadrate(?), frontal, parietal,
posterior mandible(?), atlas(?), axis (10 mm), two caudal vertebrae (10 mm),
partial clavicle(?) (49 mm), distal scapula(?) (~37 mm), ilia(?) (43 mm), partial
pubis(?) (37 mm), ischia(?) (22 mm)
Drepanosauridae- cervical vertebra (18 mm)
Pterosauromorpha- ischium
Theropoda- basioccipital, prootic, exoccipital, epiotic, supraoccipital
Coelophysoidea- (juvenile?) incomplete femur (~58 mm)
Paratype- (TTU P 9201)
unassigned- maxilla, nasal(?), lacrimal, jugal, quadratojugal, quadrate(?),
parietal(?), basioccipital(?), vomer, palatine, pterygoid, partial dentary,
posterior dentary(?), posterior mandible, axis (8 mm), cervical rib, seven dorsal
vertebrae (7-10 mm), two dorsal ribs, two sacral vertebrae (7 mm), eighteen
caudal vertebrae (7-9 mm), two chevrons, scapula(?) (~21 mm), coracoid(?) (18
mm), humerus (~39 mm), ilium(?) (22 mm), proximal tibia, proximal fibulae, metatarsal
I(?) (9 mm), phalanx (10 mm), phalanges, unguals
Drepanosauridae- seven cervical vertebrae (9-12 mm)
Pterosauromorpha- tibia (~39 mm), fibula (~41 mm), four tarsals, metatarsal
I (12 mm), metatarsal II (20 mm), metatarsal III (18 mm), metatarsal IV (14
mm)
Theropoda- distal tarsal IV, metatarsal II (25 mm), metatarsal III (27 mm),
metatarsal IV (25 mm)
Coelophysoidea- (juvenile?) astragalus, calcaneum
Late Carnian, Late Triassic
Tecovas Formation of the Dockum Group, Texas, US
Referred- (TTU P 9350-9355) six dorsal vertebrae (6-10 mm) (Chatterjee,
1995)
(TTU P 9356-9359) four caudal vertebrae (8-9 mm) (Chatterjee, 1995)
(TTU P 9360) coracoid(?) (14 mm) (Chatterjee, 1995)
(TTU P 9361) sternum(?) (25 mm) (Chatterjee, 1995)
(TTU P 9362) humerus (~80 mm) (Chatterjee, 1995)
(TTU P 9263) humerus (Chatterjee, 1995)
(TTU P 9364) partial mandible (Chatterjee, 1995)
(TTU P 9365) humerus (Chatterjee, 1995)
(TTU P 9367) pterosauromorph tibia (Chatterjee, 1995)
(TTU P 9368) pterosauromorph tibia (~58 mm) (Chatterjee, 1995)
(TTU P 9369) pterosauromorph fibula (Chatterjee, 1995)
(TTU P 9370) femur (~58 mm) (Chatterjee, 1995)
(TTU P 9371) femur (Chatterjee, 1995)
(TTU P 9372) femur (Chatterjee, 1995)
(TTU P 9373) femur (Chatterjee, 1995)
(TTU P 9374) tibia (Chatterjee, 1995)
(TTU P 9375-9380) phalanges (Chatterjee, 1995)
Diagnosis- A chimaerical taxon whose holotype includes portions of multiple
organisms.
Comments- Initially identified as a juvenile Coelophysis (Chatterjee,
1986), this material was then reported as the oldest bird (Wilford, 1986; Chatterjee,
1987, 1988). This was met with skepticism from the paleontological community,
with Ostrom (1987) questioning its avialan identification, while Paul (1988)
tentatively believed it to be a herrerasaurian.
After its official description was published (Chatterjee, 1991), some agreed
it was avialan (Kurochkin, 1992; Peters, 1994; Kurochkin, 1995; Martin, 2004).
When analyzed as a bird, Protoavis is resolved as- an ornithurine intermediate
between Avimimus and Ornithothoraces (Chatterjee, 1991; Chatterjee, 1998;
Dyke and Thorley, 1998); or an enantiornithine-grade ornithothoracine (Chatterjee,
1999). Kurochkin (1995) hypothesized it was the basalmost ornithurine (sensu
Feduccia, Martin, etc.).
However, most dismissed an avialan identity (Ostrom, 1991; Wellnhofer, 1992;
Chiappe, 1995; Feduccia, 1996; Ostrom, 1996; Sereno, 1997; Hunt et al., 1998;
Renesto, 2000; Nesbitt et al., 2005). These authors have usually suggested the
remains are chimaerical, including a non-avian coelurosaur braincase (Witmer,
2001), megalancosaurid cervicals (Renesto, 2000), possibly lepidosauromorph
humerus (Witmer, 2001), coelophysoid femur and proximal tarsals (Hutchinson,
2001; Nesbitt et al., 2005), and non-avian archosaurian pes (Sereno, 1997).
Chinle and Dockum megalancosaurids (Dolabrosaurus; Harris and Downs,
2002), lepidosauromorphs (Clevosaurus), coelophysoids (Coelophysis,
Gojirasaurus) and crurotarsans (parasuchians, aetosaurs, poposaurids,
rauisuchids) are known, but if some elements are coelurosaurian, it would be
unprecedented.
Witmer (2001) has undertaken the most detailed independant analysis of the remains,
and concludes many of the morphologies identified by Chatterjee (e.g. pterygoid
cotyla on quadrate; mandibular condyles on quadrate; quadrate foramina; quadratojugal
cotyle on quadrate; basisphenoid; scapulae; clavicle; coracoidal sulci on sternum;
intermuscular line on sternum; ulnar quill knobs; metacarpal quill knobs; opisthopuby;
fused ilium and ischium; ischial antitrochanter; tibiotarsal fusion) cannot
be confirmed. However, he notes several characters are birdlike- apparently
absent contact between squamosal and quadratojugal and postorbital; heterocoelous
cervical vertebrae; well developed cervical hypapophyses; large cervical neural
canals; coracoid morphology (elongated; strut-like; procoracoid and acrocoracoid
processes).
Witmer described the large floccular fossa, cranial pneumatic recesses and metotic
strut as coelurosaurian characters in the holotype braincase. However, the metotic
strut is a tetanurine character, the anterior and dorsal tympanic recesses are
theropod characters, the posterior tympanic recess has been described in Ceratosaurus,
and large floccular fossae are known at least as basally as Ceratosauria. Furthermore,
Protoavis' floccular fossa appears to be narrow like non-maniraptorans,
and its posterior temporal recess opens outside the otic recess unlike avialans.
So while it's possible the braincase belongs to a tetanurine or ceratosaur,
more information on coelophysoid braincase variation and ontogeny would be ideal.
Drepanosaurids share the avian cervical characters noted above with Protoavis
and the vertebrae of the two taxa are extremely similar, making it highly likely
some material is referrable to Dolabrosaurus or another drepanosaurid.
Hutchinson (2001) noted the proximal femur is identical to coelophysid femora
and suggested it may be a juvenile. Witmer (2001) found the proximal tarsals
to resemble non-coelurosaurian theropods, while Nesbitt et al. (2005) confirmed
all these elements are coelophysoid. Nesbitt et al. (2007) noted the following
dinosaur and theropod characters in the femur- offset head; ligament sulcus;
trochanteric fossa; trochanteric shelf; small posterior trochanter. They agreed
with Hutchinson it's similar to coelophysoids, but found no apomorphies of that
clade, and noted the fourth trochanter is not absent (contra Chatterjee, 1999),
but that the area is unpreserved. Nesbitt et al. also noted coelophysoid-like
tarsal characters, such a a fibular facet formed from both proximal tarsals
(unlike neoceratosaurs and tetanurines) and the distally (not anteriorly) articulated
proximal tarsals (unlike tetanurines). It's probable these belong to juvenile
Coelophysis (as originally believed) or Gojirasaurus. Additional
Coelophysis-like material is known from the Dockum Group (UCMP online;
Lehman and Chatterjee, 2005). The astragalus' low ascending process indicates
it is a coelophysoid more closely related to Coelophysis than Dilophosaurus.
Many elements of Protoavis appear to be from a new supposed pterosauromorph
named and described in a thesis (Atanassov, 2001, 2002). The original (Chatterjee,
1991; reidentified as a vomer and pterygoid in 1999) sternum is an ischium,
the radius and ulna are the tibia and fibula, and the metacarpus is the metatarsus.
The new form's maxilla, dentary, ilium and femur seem unrepresented in Protoavis'
material, while some characters of the vertebrae are a good match. Unfortunately,
only two vertebrae (both sacrals) were associated with the appendicular and
cranial elements of this new taxon, so it's possible the latter's vertebrae
are from a drepanosaurid or other taxon.
References- Chatterjee, 1986. The Late Triassic Dockum vertebrates: Their
stratographic and paleobiogeographic significance. In Padian (ed.). The beginning
of the age of dinosaurs: Faunal change across the Triassic-Jurassic boundary.
Cambridge University Press. 378 pp.
Wilford, 1986. Texas fossil may be bird's oldest ancestor. New York Times. 8-14-1986.
Chatterjee, 1987. Skull of Protoavis and early evolution of birds. JVP.
7(3) 14A.
Ostrom, 1987. Protoavis, a Triassic bird?. Archaeopteryx. 5, 113-114.
Chatterjee, 1988. Functional significance of the semilunate carpal in archosaurs
and birds. JVP. 8(3) 11A.
Paul, 1988. Predatory dinosaurs of the world. Simon and Schuster, New York.
A New York Academy of Sciences Book. 464 pp.
Ostrom, 1991. The bird in the bush. Nature. 353, 212.
Chatterjee, 1991. Cranial Anatomy and relationships of a new Triassic bird from
Texas. Philosophical Transactions of the Royal Society of London Series B 332(1265):
277-342.
Kurochkin, 1991. Protoavis, Ambiortus and other paleornithological
rarities. [in Russian]. Priroda. 1991, 43-53.
Chatterjee, 1992. Texas fossil identified as the world's oldest known bird.
J. Geol. Soc. India 39 p. 89-90.
Viohl, 1992. No evidence of Triassic birds. [in German]. Archaeopteryx. 10,
77-79.
Wellnhofer, 1992. Protoavis: The oldest bird?. [in German]. Naturwiss.
Rundsch. 45, 107-108.
Kurochkin, 1992. The oldest bird. [in Russian]. Priroda. 1992, 100-101.
Peters, 1994. Die Entstehung der Vogel-Verandern die jungsten Fossilfunde das
Modell? In Gutmann et al. (eds.). Morphologie und evolution. Frankfurt. 403-424.
Chatterjee, 1995. The Triassic bird Protoavis. Archaeopteryx. 13, 15-31.
Chiappe, 1995. The first 85 million years of avian evolution. Nature. 378, 349-355.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of Class Aves.
Archaeopteryx. 13, 47-66.
Feduccia, 1996. The Origin and Evolution of Birds. Yale University Press. 420
pp.
Ostrom, 1996. The Questionable Validity of Protoavis. Archaeopteryx.
14, 39-42.
Chatterjee, 1997. The Rise of Birds: 225 Million Years of Evolution. John Hopkins
University Press. 312 pp.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth
and Planetary Sciences. 25, 435-489.
Chatterjee, 1998. The avian status of Protoavis. Archaeopteryx. 16, 99-122.
Dyke and Thorley, 1998. Reduced cladistic consensus methods and the inter-relationships
of Protoavis, Avimimus, and Mesozoic birds. Archaeopteryx. 16,
123-129.
Hunt, A.P., S.G. Lucas, A.B. Heckert, R.M. Sullivan and M.G. Lockley. 1998.
Late Triassic dinosaurs from the western United States. Geobios, 31(4):511-531.
Chatterjee, 1999. Protoavis and the early evolution of birds. Palaeontographica
A. 254, 1-100.
Renesto, 2000. Bird-like head on a chameleon body: new specimens ofthe enigmatic
diapsid reptile Megalancosaurus from the Late Triassic of northern Italy.
Rivista Italiana di Paleontologia e Stratigrafia. 106:157180.
Atanassov, 2001. Two new archosauromorphs from the Late Triassic of Texas. Journal
of Vertebrate Paleontology. 21(3) 30A.
Hutchinson, 2001. The evolution of femoral osteology and soft tissues on the
line to extant birds (Neornithes). Zoological Journal of the Linnaean Society.
131, 169-197.
Witmer, 2001. The role of Protoavis in the debate on avian origins. In
Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution
of Birds. Yale University. 538-548.
Atanassov, 2002. Unpublished thesis. Texas Tech University.
Harris and Downs, 2002. A drepanosaurid pectoral girdle from the Ghost Ranch
(Whitaker) Coelophysis Quarry (Chinle Group, Rock Point Formation, Rhaetian),
New Mexico. JVP. 22(1), 70-75.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica.
50(6), 978-990.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur
record, part 3: Saurischians of North America. JVP 25(3) 96A.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
Liliensternus Welles,
1984
Diagnosis- (after Rauhut, 2000) broad rounded ridge on the cervical vertebrae,
that extends from the posterior end of the diapophyses to the posteroventral
end of the vertebral centrum.
L. liliensterni (Huene, 1934) Welles, 1984
= Halticosaurus liliensterni Huene, 1934
Norian, Late Triassic
Trossingen Formation, Germany
Syntypes- (MB.R.2175; = HMN R1291) (gracile young subadult) (5.15 m,
127 kg) (partial skull 395 mm), ventral maxilla, jugal, postorbital, incomplete
squamosal, quadrate, dentary, surangular, partial angular, fourth cervical vertebra
(70 mm), tenth cervical vertebra (47 mm), first sacral vertebra (52 mm), second
sacral vertebra (52 mm), third sacral vertebra (52 mm), scapula (~240 mm), humerus
(214 mm), radius (151 mm), ulna (158 mm), metacarpal II (70 mm), ilium (270
mm), pubis (418 mm), ischium (303 mm), femur (440 mm), tibia (409 mm), astragalocalcaneum
(79 mm wide, astragalus 60 mm wide), metatarsal II (214 mm), phalanx II-1 (73
mm), phalanx II-2 (39 mm), pedal ungual II (55 mm), metatarsal III (230 mm),
phalanx III-1 (69 mm), phalanx III-2 (55 mm), phalanx III-3 (46 mm), pedal ungual
III (46 mm), metatarsal IV (221 mm), phalanx IV-1 (51 mm), phalanx IV-2 (35
mm), phalanx IV-3 (29 mm), phalanx IV-4 (27 mm)
(MB.R.2175) (gracile young subadult) partial skeleton
Diagnosis- (after Rauhut, 2000) one pair of pleurocoels in the cervical
vertebrae, less developed infradiapophyseal fossa; the absence of a horizontal
ridge at the basis of the cervical neural spines; absence of a lateral bulge
on the ilium.
Comments- The syntype remains of this species are usually referred to
two individuals. However, the material was found disarticulated and may represent
more than two individuals. Because of this and the fact it is hard to separate
the remains belonging to the various individuals, Rauhut and Hungerbuhler (2000)
recommend retaining all of the material as the syntypes of Liliensternus
(contra Welles 1984, who made the larger individual the syntype). Because the
neurocentral sutures are unfused and only two fused sacrals are present, the
remains are probably juveniles or subadults. Contra Rowe and Gauthier (1990),
the pubis encloses a complete obturator foramen, not just a notch.
References- Huene, 1934. Ein neuer Coelurosaurier in der thuringischen
Trias. Pal. Zeit. 16 145-170, 4 pls.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie 185 p. 85-180.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia
15, 75-88.
L? sp. indet. (Sander, 1992)
Norian, Late Triassic
Trossingen Formation, Germany
Material- (UA coll.) proximal metatarsal II (Galton, 2000)
at least two occurances (Sander, 1992)
Comments- The UA coll. metatarsal was originally a syntype of Plateosaurus
engelhardti, described as a manual or pedal element by Meyer (1855) and
a pubic fragment by Huene (1908). It was reidentified as a proximal metatarsal
IV by Galton (2000) and a proximal metatarsal II by Moser (2003). Comparison
with figures of Liliensternus indicates Moser is correct.
Reference- Meyer, 1955. Zur fauna der Vorwelt. Die saurier des Muschelkalkes,
rnit Rucksicht auf die saurier aus Buntem Sandstein und Keuper. 167 pp.
Huene, 1908. Die Dinosaurier der Europaiaschen Triasformation mit Berucksichtiging
der aussereuropaischen Vorkommnisse. Geol. Paleont. Abhandl. Suppl. 1, 1-419.
Sander, 1992. The Norian Plateosaurus bonebeds of central Europe and
their taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology. 93:255-296.
Galton, 2000. The prosauropod dinosaur Plateosaurus Meyer, 1837 (Saurischia:
Sauropodomorpha). I. The syntypes of P. engelhardti Meyer, 1837 (Upper
Triassic, Germany), with notes on other European prosauropods with "distally
straight" femora. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen.
216(2): 233-275.
Moser, 2003. Plateosaurus engelhardti Meyer, 1837 (Dinosauria: Sauropodomorpha)
aus dem Feuerletten (Mittelkeuper; Obertrias) von Bayern. Zitteliana B 24, 3-186.
L? sp. indet. (Sander, 1992)
Norian, Late Triassic
Knollenmergel, Switzerland
Reference- Sander, 1992. The Norian Plateosaurus bonebeds of central
Europe and their taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology.
93:255-296.
unnamed clade (Zupaysaurus rougieri + Lophostropheus airelensis
+ Coelophysis bauri)
Diagnosis- (modified from Tykoski, 2005) astragalus and tibia fused in
adults.
(after Ezcurra and Novas, 2006)- angle between the rostrodorsal margin of the
maxilla and the alveolar margin less than 75 degrees (unknown in Lophostropheus);
infratemporal fenestra strongly rostrocaudally compressed, with maximum length
versus maximum length of the orbit ratio less than 0.8 (unknown in Lophostropheus);
caudal curvature of the dorsal end of quadrate, with quadrate head caudodorsally
oriented (unknown in Lophostropheus).
(after Ezcurra and Cuny, 2007) supraacetabular crest continuous with the lateral
margin of the brevis fossa, forming a well-developed ridge, without a notch
between both structures (unknown in Zupaysaurus).
Lophostropheus Ezcurra
and Cuny, 2007
L. airelensis (Cuny and Galton, 1993) Ezcurra and Cuny, 2007
= Liliensternus airelensis Cuny and Galton, 1993
Late Rhaetian-Early Hettangian, Late Triassic-Early Jurassic
Moon-Airel Formation, France
Holotype- (Caen University coll.) tooth, five cervical vertebrae (72,
83 mm), two posterior dorsal vertebrae (75 mm), four sacral vertebrae, several
caudal vertebrae (72 mm), partial ilium, proximal pubes, partial ischium
Diagnosis- (after Rauhut, 2000) deep infradiapophyseal fossa in anterior
cervical vertebrae; horizontal ridge at the basis of the neural spine in cervical
vertebrae; ilium with a triangular lateral bulge above the supraacetabular crest.
(after Ezcurra and Cuny, 2007) moderately convex anterior articular surface
of the anterior postaxial cervical vertebrae (also in Ceratosauria+Tetanurae);
large and
oval lateral fossa on last dorsal vertebral centrum (also in Herrerasaurus);
dorsoventrally well-extended hyposphene in the last dorsal vertebra; incipient
concavity on the cranial articular surface of cranial caudal vertebrae (also
present in Ceratosauria+Tetanurae); constant length of caudal vertebrae along
the tail (also in Dilophosaurus).
Comments- Rauhut and Hungerbuhler (2000) also listed the "cervical
vertebrae with dorso-ventrally narrow, antero-posteriorly elongated posterior
pleurocoel" as being diagnostic, but it is also present in Coelophysis.
Although Rauhut and Hungerbuhler (2000) believe this is a valid species, they
think characters such as the presence of two pairs of cervical pleurocoels (versus
one pair in L. liliensterni), may indicate it deserves a separate genus.
References- Larsonneur and Lapparent, 1966. Un dinosaurien carnivore,
Halticosaurus, dans le Réthien d´ Airel (Manche). Bulletin
Societe Linneenne de Normandie. 10:108116.
Cuny and Galton, 1993. Revision of the Airel theropod dinosaur from the Triassic-Jurassic
boundary (Normandy, France). Neues Jahrbuch für Geologie und Paläontologie,
Abhandlungen 187 p. 261-288.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia
15, 75-88.
Ezcurra and Cuny, 2007. The coelophysoid Lophostropheus airelensis, gen.
nov.: a review of the systematics of "Liliensternus" airelensis
from the Triassic-Jurassic boundary outcrops of Normandy (France). 73-86.
Zupaysaurus Arcucci and Coria,
2003
= Zupaysaurus unpublished, 1999
Z. rougieri Arcucci and Coria, 2003
Late Norian-Rhaetian, Late Triassic
Upper Los Colorados Formation, Argentina
Holotype- (PULR-076) (5-6 m) skull (450 mm), mandibles, atlas, axis (110
mm), cervical vertebrae, cervical ribs, dorsal vertebrae, sacral vertebrae,
distal caudal vertebrae, proximal scapulocoracoid, two proximal manual unguals,
distal femora, proximal tibia, distal tibia, distal fibula, astragalocalcaneum
Diagnosis- (modified from Arcucci and Coria, 2003) horizontal ramus of
the maxilla with parallel dorsal and ventral margins; tibia with a very deep
and caudally open notch for the reception of an astragalar caudal process.
(after Ezcurra, 2006) maxillary fenestra within the antorbital fossa (also in
Tetanurae); ventrally bowed rostral process of the lacrimal (also in Sinraptor);
kinked ventral process of the squamosal; wide contact between squamosal and
quadratojugal (also in several tetanurines).
(after Ezcurra and Novas, 2006) maxillary-jugal ventral margin describing an
obtuse angle in lateral view; notch on the dorsal margin of the ascending process
of the maxilla, relating to horizontal ramus of lacrimal rostrally tapering
onto the forked caudal tip of the ascending process of the maxilla; lacrimal
with highly pneumatized antorbital recess (also in Ceratosauria+Tetanurae);
short and square-shaped retroarticular process of the mandible; cnemial crest
poorly developed.
Comments- Originally thought to be "more derived than Coelophysis"
(Arcucci and Coria, 1997), and later a ceratosaur (Arcucci and Coria, 1998).
When officially described by Arcucci and Coria (2003), it was found to be a
basal tetanurine. However, later analyses have found that it is in fact a coelophysoid.
Carrano et al. (2005) found it to be a coelophysoid, but could not recover any
resolution within that clade. Zupaysaurus was found to be in a trichotomy
with Liliensternus and coelophysids by Tykoski (2005). Ezcurra and Novas
(2005) agree the taxon is a coelophysoid, which was elaborated on in Ezcurra
and Novas (2006). Of the supposed tetanurine characters, some are reinterpreted
as convergences (maxillary fenestra; lacrimal recess; fibula with distal end
expanded almost double the shaft width), while others aren't present in Zupaysaurus
(antorbital maxillary tooth row; lacrimal horn; distal tibia transversely expanded)
or are present in coelophysoids as well (lateral temporal fenestra reduced and
key-hole-shaped; tibia with a caudolaterally concave distal end; ascending process
of astragalus cranially positioned). They found it to be in a trichotomy with
Segisaurus and Coelophysidae. Smith et al. (2007) found Zupaysaurus
to be more derived than coelophysoids, sister to a clade containing dilophosaurids,
ceratosaurs and tetanurines. This was based on- tooth row that ends at the anterior
rim of the orbit (not actually present); jugal with an expanded anterior end;
lacrimal fenestra; broad contact between the squamosal and quadratojugal; well-developed
anterior wall to the lateral mandibular glenoid; broadened retroarticular process;
posterodorsally facing surface for the attachment of the m. depressor mandibulae
on the retroarticular process; astragalar ascending process that is higher than
the astragalar body. Placing Zupaysaurus inside Coelophysoidea took five
more steps.
References- Arcucci and Coria, 1997. First record of Theropoda (Dinosauria
- Saurischia) from the Los Colorados Formation (Upper Triassic, La Rioja, Argentina).
Ameghiniana 34(4) 531.
Arcucci and Coria, 1998. Skull features of a new primitive theropod from Argentina.
JVP 18(3) 24A-25A.
unpublished, 1999. Discovery News. 11-12-1999.
Arcucci and Coria, 2003. A new Triassic carnivorous dinosaur from Argentina.
Ameghiniana 40(2): 217 - 228.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli,
a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate
Paleontology. 25(4), 835849.
Ezcurra and Novas, 2005. Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NW Argentina. In Kellner, Henriques and Rodrigues
(eds.), II Congresso Latino-Americano de Paleontologia de Vertebrados, Boletim
de Resumos. Museum Nacional/UFRJ, Rio de Janeiro 102-104.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. The cranial anatomy of the coelophysoid theropod Zupaysaurus
rougieri from the Upper Triassic of Argentina. Historical Biology. iFirst
Article, 18 pp. DOI 10.1080/08912960600861467.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article,
38 pp. DOI 10.1080/08912960600845791.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti
(Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications
for early theropod evolution. Zoological Journal of the Linnean Society. 151,
377-421.
unnamed coelophysoid (Heckert, Zeigler, Lucas, Rinehart and Harris,
2000)
Norian, Late Triassic
Petrified Forest Formation of Chinle Group, New Mexico
Material- (NMMNH P-29046) femora, tibia, proximal tibia, proximal fibula
(NMMNH P-29047) partial ilium, incomplete ischium, proximal tibia
(NMMNH P-30779) dorsal vertebra
(NMMNH P-30780) dorsal vertebra
(NMMNH P-30852) premaxilla, maxilla, lacrimal, dentaries, splenial, surangulars,
hyoids, two anterior cervical vertebrae, cervical ribs
(NMMNH P-31293) incomplete tibia
(NMMNH P-31661) sacrum, partial scapulocoracoid
Diagnosis- appressed lateral surface of tibial shaft.
Comments- Originally referred to Eucoelophysis sp. by Heckert
et al. (2000) based on supposed similarities in the scapulocoracoid, ischium
and tibia, Nesbitt et al. (2007) showed the former genus is non-dinosaurian
while Heckert et al.'s material exhibits numerous dinosaur, theropod and coelophysoid
synapomorphies. Eucoelophysis' holotype differs in lacking a distinct
acetabular rim on the ischium, an offset femoral head, a differentiated lateral
and fibular condyle that is separated by a distinct sulcus, and a well developed
curved cnemial crest. The appressed lateral tibial surface is developed differently
in both taxa. Ezcurra (2006) noted NMMNH P-30852 shows an alveolar ridge like
Liliensternus and more derived coelophysoids, and a square anterior end
on the antorbital fossa as in Zupaysaurus and more derived coelophysoids.
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, 2732.
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, 127132.
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), 209243.
Coelophysidae Nopcsa, 1928 sensu Welles,
1984
Definition- (Coelophysis bauri + Megapnosaurus rhodesiensis
+ "Megapnosaurus" kayentakatae) (Holtz, 1994)
Other definitions- (Coelophysis bauri + Procompsognathus triassicus)
(Sereno, in press; modified from Sereno, 1998)
(Coelophysis bauri + Megapnosaurus rhodesiensis) (modified from
Tykoski and Rowe, 2004)
= Podokesauridae Huene, 1914
= Podokesaurinae Huene, 1914 sensu Nopcsa, 1923
= Segisauridae Camp, 1936
Diagnosis- (modified from Tykoski, 2005) medial flange of fibula overlaps
astragalar ascending process; fibular facet of astragalus reduced, subtriangular
in shape on anterolateral corner of proximal surface; postaxial cervical neural
arches house pneumatic cavities lateral to neural canal; lateral surface of
postaxial cervical neural arch pediucels have posteriorly directed fossae anterior
to postzygapophyses; elongate mid cervical centra (>3 times height of anterior
articular surface); elongate posterior dorsal centra (>2 times height of
anterior articular surface); m. iliofemoralis fossa of ilium stops short of
posterior margin, leaving distinct lateral rim on postacetabular process; distal
tarsal III fuses to metatarsal III in adults; metatarsals II and III fuse in
adults; circular orbit; few or no neurovascular foramina on lateral premaxillary
surface; medial surface of anteromedial maxillary process smooth; low rounded
horizontal ridge on lateral surface of jugal; ventral process of squamosal narrow;
lamina spanning the basal tubera bears small median spur that projects anteriorly
along roof of basisphenoid recess; anterior premaxillary teeth subcircular in
section; anterior premaxillary teeth with reduced curvature; cervical ribs elongate
(>4 times central length); posterior margin of ilium concave in lateral view.
Comments- Tykoski (2005) also included the presence of a furcula as an
ambiguous synapomorphy of this clade, possibly based on a miscoding of Carnotaurus.
Podokesauridae- Podokesauridae was first proposed by Huene (1914) as
a basal grade of coelurosaurs (sensu Huene). This position was retained throughout
the mid-1900's, with podokesaurids seen as directly ancestral to coelurids,
and less directly to compsognathids and ornithomimids. Barsbold (1977) had podokesaurids
ancestral to all theropods, and coelurids and troodontids in particular. Huene
(1914) originally included Podokesaurus, Coelophysis, Saltopus
and Tanystropheus. Later added were Procompsognathus (Huene, 1914),
Halticosaurus (Huene, 1914), Procerosaurus (Huene, 1920), Avipes
(Huene, 1932), Spinosuchus (Huene, 1932), Dolichosuchus (Romer,
1956), Lukousaurus (Romer, 1956), Velocipes (Romer, 1956), Trialestes
(Reig, 1963; as Triassolestes), Megapnosaurus (Raath, 1969; as
Syntarsus), Dilophosaurus (Russell, 1984), Alwalkeria (Chatterjee,
1987; as Walkeria), Segisaurus (Carroll, 1988), "Comanchesaurus"
(Hunt and Lucas, 1989), Sarcosaurus (Madsen and Welles, 2000) and Liliensternus
(earlier as Halticosaurus liliensterni, then explicitly by Madsen and
Welles, 2000). The family was generally not used after the 1980's, with its
genera then referred to as ceratosaurs, coelophysoids and/or coelophysids. Madsen
and Welles (2000) correctly noted that Podokesauridae has priority over Coelophysidae
according to the ICZN, since it was named fourteen years earlier. Yet all other
workers ignore this, generally because they value Phylocode rules and Podokesauridae
has not been phylogenetically defined, or they misread the ICZN to demand family-level
eponyms be diagnosable (it actually only says they must be valid). Coelophysidae
is only used on this website because of its near universal usage since 1991.
Nopcsa (1923) changed it to subfamily Podokesaurinae within Hallopodidae, which
has not been followed as Hallopus is a crocodylomorph.
Coelophysidae- Welles (1984) erected Coelophysidae from Nopcsa's earlier
subfamily Coelophysinae, including Coelophysis and Sarcosaurus
without comment. Paul (1988) later included Coelophysis, Dilophosaurus,
Elaphrosaurus, Liliensternus and Megapnosaurus (as Coelophysis
rhodesiensis) in the family. This was followed by Novas (1991, 1992), though
Holtz (1994) restricted the term to exclude the basal Dilophosaurus.
This is followed by all workers currently yet the precise extent of Coelophysidae
has since been controversial, sometimes including Liliensternus (Carrano
et al., 2002), and sometimes restricted to Coelophysis and Megapnosaurus
(Rauhut, 2003). This depends both on the definition used and the resolution
within derived Coelophysoidea. An intermediate extent is used here, based on
the definition in Holtz (1994) and the analysis of Tykoski (2005).
Segisauridae- Segisauridae was established by Camp (1936) to include
only Segisaurus, viewed as related to coelurids and compsognathids within
Coelurosauria sensu Huene. This practice was followed until Gauthier (1986)
determined that Segisaurus was a ceratosaur sensu lato. Segisauridae
has been almost unused since, with Segisaurus being placed in Coelophysoidea
and/or Coelophysidae instead (Sereno, 1997). If Coelophysidae is restricted
to Coelophysis + Megapnosaurus, as defined in Tykoski and Rowe
(2004), then Segisauridae may be used for its sister clade, including Segisaurus,
"Megapnosaurus" kayentakatae and perhaps Camposaurus.
On this site they are grouped as Segisaurinae within Coelophysidae.
Ex-coelophysids- Huene (1914) included Saltopus as one of his
original podokesaurids, placing it ancestral to Halticosaurus. This placement
was common, though current research indicates it is most likely a more basal
dinosauriform (Langer, 2004). Tanystropheus is another original podokesaurid
of Huene's, but is now known to be a protorosaur (Wild, 1973). Several species
(T. bauri, T. longicollis and T. willistoni) are now recognized
as coelophysids however. Halticosaurus was assigned to Podokesauridae
(Huene, 1914) by many authors and indeed seems to be coelophysoid, though its
short cervical centra exclude it from Coelophysidae as used here. Procompsognathus
was soon placed in Podokesauridae by Huene (1914), which has been supported
by some recent analyses (Ezcurra and Novas, 2006) and rejected by others (Allen,
2004). Procerosaurus was assigned to Podokesauridae by Huene (1920),
but is actually a junior synonym of Tanystropheus (Wild, 1973). Spinosuchus
was first described as a Coelophysis specimen (Case, 1922), and was later
referred to Podokesauridae once it was placed in its own genus by Huene (1932).
Nesbitt et al. (2007) have most recently assigned it to Archosauriformes incertae
sedis. Avipes was first described as a podokesaurid (Huene, 1932),
but recent studies have been unable to place it more definitively than Avemetatarsalia
(Rauhut and Hungerbuhler, 2000). Lukousaurus was originally described
as a coelurosaur related to Podokesaurus, Procompsognathus and
Saltopus (Young, 1948), and explicitly placed in Podokesauridae by Romer
(1956). It has not been recently restudied, but lacks several coelophysoid apomorphies
and is more likely crurotarsan. Velocipes was placed in Podokesauridae
by Romer (1956), yet like Avipes, recent studies have been unable to
place it more definitively than Avemetatarsalia (Rauhut and Hungerbuhler, 2000).
Dolichosuchus was similarly placed in Podokesauridae by Romer (1956),
but Welles (1984) notes it is almost identical to Liliensternus, so it
is possibly just outside Coelophysidae. Reig (1963) assigned Trialestes
(as Triassolestes) to Podokesauridae. Romer (1972) properly recognized
it as a crurotarsan, though it does have several dinosaur-like characters. Russell
(1984) assigned Dilophosaurus to the Podokesauridae, though following
Holtz (1994), it is now considered just basal to Coelophysidae. Welles (1984)
placed Sarcosaurus in Coelophysidae without comment, while Madsen and
Welles (2000) placed it in Podokesauridae, but it appears to represent a more
basal coelophysoid (Tykoski, 2005). Protoavis was originally believed
to be a juvenile Coelophysis (Chatterjee, 1986), and while some material
is coelophysoid (Nesbitt et al., 2007), none can be assigned to Coelophysidae
itself with certainty. (Chatterjee (1987) assigned his new genus Walkeria
to Podokesauridae, but it has since been renamed Alwalkeria and placed
more basally as a basal eusaurichian (Langer, 2004). Elaphrosaurus was
assigned to Coelophysidae by a couple authors (Paul, 1988; Novas, 1992) before
it was realized to be a ceratosaur sensu lato (Holtz, 1994), which has been
the result of every published cladistic analysis. Liliensternus is often
assigned to Coelophysidae (e.g. Paul, 1988) and was often viewed as a podokesaurid
when still classified as a species of Halticosaurus, yet based on the
present definition of Coelophysidae is just barely excluded from that clade.
"Comanchesaurus" was listed as a podokesaurid by Hunt and Lucas (1989),
though it more recently could not be definitely placed closer to Coelophysis
than Liliensternus (Nesbitt et al., 2007). Sullivan and Lucas (1999)
believed their new taxon Eucoelophysis to be most closely related to
Coelophysis and Megapnosaurus, but Ezcurra (2006) has since determined
it to be a more basal dinosauriform. Rauhut (2003) found Shuvosaurus
to fall out as a coelophysid in his analysis, sister to Megapnosaurus,
but the description of Effigia (Nesbitt and Norell, 2006) led to its
recognition as a crurotarsan.
Defining Coelophysidae- Holtz (1994) first defined Coelophysidae, as
the most exclusive clade containing Coelophysis bauri, Syntarsus rhodesiensis
and S. kayentakatae. This matches its use in recent papers such as Tykoski
(2005) and Ezcurra and Novas (2006). Two other definitions have been proposed
for Coelophysidae, neither one ideal. Sereno (1998) used Procompsognathus
and Coelophysis as internal specifiers. Yet Procompsognathus may
not be dinosaurian (Allen, 2004), in which case all of Dinosauria would be coelophysids
according to his definition. Even when Procompsognathus is found to be
a coelophysoid, it usually has an uncertain position within the clade. Tykoski
(2005) found it could fall it anywhere in the Liliensternus + Coelophysis
clade. Ezcurra and Novas (2006) found it only slightly more constrained, within
the Zupaysaurus + Coelophysis clade. Under this definition, Coelophysidae
has an extremely uncertain content, which may include Liliensternus,
Zupaysaurus, Segisaurus, etc. or may not. Tykoski and Rowe (2004)
proposed a definition which used Coelophysis and Megapnosaurus rhodesiensis
as internal specifiers. This is easy to apply to most analyses, where the two
emerge as sister taxa. However, this would exclude not only Liliensternus,
Zupaysaurus and Lophostropheus from Coelophysidae, but also Segisaurus
and "Megapnosaurus" kayentakatae. This is far more exclusive
than most concepts of Coelophysidae (or Podokesauridae) have been. Also, the
synapomorphies for Coelophysis + Megapnosaurus are all cranial,
making the assignment of postcrania to Coelophysidae impossible.
References- Huene, 1914. Beiträge zur geschichte der Archosaurier
[Contribution to the history of the archosaurs]. Geologie und Paläontologie
Abhandlungen. 13(7), 1-56.
Huene, 1914. The dinosaurs not a natural order. The American Journal of Science,
series 4. 38, 145-146.
Huene, 1914. Das natürliche System der Saurischia [The systematics of the
Saurischia]. Centralblatt für Mineralogie, Geologie und Paläontologie.
1914,154-158.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien
[Phylogenetic results of some investigations of Triassic reptiles]. Zeitschrift
für Induktive Abstammungsund Vererbungslehre. 24, 159-163.
Case, 1922. New reptiles and stegocephalians from the Upper Triassic of Western
Texas. Carnegie Institution of Washington Publication. 321, 1-84.
Nopcsa, 1923. Die Familien der Reptilien [The families of reptiles]. Forschritte
der Geologie und Palaeontologie. 2, 1-210.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Huene, 1929. Kurze Übersicht über die Saurischia und ihre natürlichen
Zusammenhänge [A brief survey of the Saurischia and their natural context].
Paläontologische Zeitschrift. 11, 269-273.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monog. Geol. Pal. 1, pts 1 and 2, V111 + 361 pp.
Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of
Arizona. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 24 39-56, 8 figs., 2 pls.
Young, 1948. On two new saurischians from Lufeng, China. Bulletin of the Geological
Society of China. 28(1-2), 78-90.
Romer, 1956. Osteology of the Reptiles, University of Chicago Press. 1-772.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press,
Chicago. 1-468.
Raath, 1969. A new coelurosaurian dinosaur from the Forest Sandstone of Rhodesia.
Arnoldia (Rhodesia). 4(28), 1-254.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia
of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 1-87.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus
(Bassani) (Neue Ergebnisse) . Schweizerische Palaontologische Abhandlungen.
95, 1-162.
Barsbold, 1977. O evolutsiy chishcheich dinosavrov [On the evolution of carnivorous
dinosaurs]. Trudy - Sovmestnaya Sovetsko-Mongol'skaya Paleontologicheskaya Ekspeditsiya.
4, 48-56.
Russell, 1984. A check list of the families and genera of North American dinosaurs.
Syllogeus. 53, 1-35.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology
and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit.
Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Chatterjee, 1986. The Late Triassic Dockum vertebrates: Their stratographic
and paleobiogeographic significance. In Padian (ed.). The beginning of the age
of dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge
University Press. 378 pp.
Chatterjee, 1987. A new theropod dinosaur from India with remarks on the Gondwana-Laurasia
connection in the Late Triassic. Gondwana 6: Stratigraphy, Sedimentology and
Paleontology, G. D. McKenzie (ed.), Geophysical Monographs. 41, 183-189.
Carroll, 1988. Vertebrate Paleontology and Evolution. W.H. Freeman and Company.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Hunt and Lucas, 1989. Late Triassic vertebrate localities in New Mexico. pp.
72-101. in Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American
Southwest. New Mexico Museum of Natural History, Albuquerque.
Novas, 1991. Relaciones filogeneticas de los dinosaurios teropodos ceratosaurios
[Phylogenetic relationships of ceratosaurian theropod dinosaurs]. 28(3-4), 401.
Novas, 1992. La evolucion de los dinosaurios carnivoros [The evolution of carnivorous
dinosaurs]. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico:
Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez",
Cuenca, Argentina. 126-163.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth
and Planetary Sciences. 25, 435-489.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und
Paläontologie, Abhandlungen. 210(1), 41-83.
Sullivan and Lucas, 1999. Eucoelophysis baldwini, a new theropod dinosaur
from the Upper Triassic of New Mexico, and the status of the original types
of Coelophysis. Journal of Vertebrate Paleontology 19(1): 81-90.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey, 80 pages.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia.
15, 75-88.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri,
a small abelisauroid (Dinosauria:Theropoda) from the Late Cretaceous of Madagascar.
Journal of Vertebrate Palaeontology. 22, 510534.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Allen, 2004. The phylogenetic status of Procompsognathus revisited. JVP
24(3).
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria
Second Edition. University of California Press. 861 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur
Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic
of New Mexico, USA. Geodiversitas. 28 (4),649-684.
Nesbitt and Norell, 2006. Extreme convergence in the body plans of an early
suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of
the Royal Society B. 273, 1045-1048.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
unnamed Coelophysidae (Cope, 1887)
Norian, Late Triassic
Petrified Forest or Rock Point Formation of the Chinle Group, New Mexico, US
Material- ?(AMNH 2702; syntype of Coelurus longicollis) mid caudal
vertebra (Cope, 1887)
?(AMNH 2703) manual ungual (Cope, 1887)
(AMNH 2704; syntype of Coelurus longicollis) femur (Cope, 1887)
(AMNH 2706) pubis (226 mm) (Cope, 1887)
?(AMNH 2707) manual phalanx I-1 (Cope, 1887)
(AMNH 2708; not lectotype of Coelophysis bauri, contra Welles, 1984)
ilium (~155 mm) (Cope, 1887)
?(AMNH 2715; syntype of Coelurus longicollis) dorsal centrum (Cope, 1887)
?(AMNH 2716) ischium(?) (Cope, 1887)
(AMNH 2717; syntype of Coelurus bauri) third or fourth cervical centrum
(53 mm) (Cope, 1887)
(AMNH 2718) distal ischium (Cope, 1887)
?(AMNH 2719) proximal ischium(?) (Cope, 1887)
(AMNH 2720) partial fourth or fifth cervical vertebra (Cope, 1887)
(AMNH 2721) proximal metatarsal III (Cope, 1887)
(AMNH 2722; lectotype of Coelophysis bauri) sacrum (36, 35 mm) (Cope,
1887)
?(AMNH 2723) dorsal centrum (Cope, 1887)
(AMNH 2724) proximal pubis (Cope, 1887)
?(AMNH 2727; syntype of Tanystropheus willistoni) distal caudal centrum
(Cope, 1887)
?(AMNH 2728) distal metatarsal IV(?) (Huene, 1915)
?(AMNH 2729) dorsal transverse process (Huene, 1915)
?(AMNH 2730) distal metatarsal III (Huene, 1906)
?(AMNH 2731) proximal pubis (Huene, 1915)
?(AMNH 2732) five vertebral fragments (Padian, 1986)
?(AMNH 2734) incomplete mid caudal vertebra (Huene, 1915)
?(AMNH 2735) partial distal caudal centrum (Cope, 1887)
?(AMNH 2736) partial cervical neural arch (Huene, 1915)
?(AMNH 2737) proximal humerus (Huene, 1915)
?(AMNH 2738) ischial fragments (Huene, 1915)
?(AMNH 2739) dorsal centrum (Huene, 1915)
?(AMNH 2740) distal metatarsal III (Huene, 1915)
?(AMNH 2742) bone fragments (Padian, 1986)
?(AMNH 2743) vertebra, six fragments (Padian, 1986)
?(AMNH 2744) partial proximal caudal vertebra (Huene, 1906)
?(AMNH 2745) proximal fibula (Huene, 1915)
?(AMNH 2746) five caudal vertebra fragments (Padian, 1986)
?(AMNH 2747) pedal fragments (Padian, 1986)
?(AMNH 2748) vertebral fragments (Padian, 1986)
?(AMNH 2749) dorsal centrum (Huene, 1915)
?(AMNH 2750) anterior sacrum (Huene, 1915)
?(AMNH 2751) posterior cervical vertebra (Huene, 1915)
?(AMNH 2752) partial anterior dorsal vertebra (Huene, 1915)
?(AMNH 2753) vertebral and bone fragments (Padian, 1986)
Comments- This material was collected with the lectotypes of Coelophysis
bauri, C. longicollis, C. willistoni and the holotype of Longosaurus.
It has been referred to various combinations of these species by different authors
(Cope, 1887, 1887; Huene, 1906, 1915; Welles, 1984; Padian, 1986). Most recently,
it was viewed as generically indeterminate (along with the types of the above
species), and Coelophysis was assigned a neotype from a different locality
(Hunt and Lucas, 1991; ICZN, 1996). Sullivan and Lucas (1999) erected a new
species of coelophysid from the Coelophysis lectotype locality, Eucoelophysis
baldwini, and referred AMNH 2706 to it. In addition, they felt the rest
of the material listed above was probably referrable to Eucoelophysis,
but could not definitively assign it due to its undiagnostic nature. Nesbitt
et al. (2005, 2007) and Ezcurra (2006) determined Eucoelophysis was not
a theropod, but a more basal dinosauriform. Nesbitt et al. (2007) reexamined
the specimens and found that the cervical vertebrae (AMNH 2717, 2720), sacrum
(AMNH 2722) and pelvic elements (AMNH 2705, 2706, 2708, 2722) could be referred
to coelophysoids based on- dual pairs of cervical pleurocoels, pubic obturator
foramen, well developed supraacetabular crest that arcs ventrally at its lateral
margin; a squared-off distal portion of the postacetabular process; a deep brevis
fossa where the lateral ridge originates near the supraacetabular crest; flattened
dorsal margin of the iliac blade; fully perforated acetabulum. It's probable
only one species of coelophysid is represented, possibly congeneric or conspecific
with C. bauri. The dorsal vertebrae and most of the limb elements were
undiagnostic within Archosauria, and a distal femur (AMNH 2725) is referrable
to a shuvosaurid.
References- Cope, 1887. The dinosaurian genus Coelurus. Amer.
Naturalist xxi pp. 367-369.
Cope, 1887. A contribution to the history of the Vertebrata of the Trias of
North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls. i, ii.
Huene, 1906. Ueber die Dinosaurier der Aussereuropaischen Trias. Geol. Pal.
Abh. 12, 99-156.
Huene, 1915. On reptiles of the New Mexican Trias in the Cope collection. Bulletin
American Museum of Natural History. 34, 485-507.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic
dinosaur from New Mexico (USA). Paläontol. Z. 65 p. 191-198.
International Commision on Zoological Nomenclature, 1996. Opinion 1842. Coelurus
bauri Cope, 1887 (currently Coelophysis bauri; Reptilia, Saurischia):
lectotype replaced by a neotype. Bulletin of Zoological Nomenclature. 53, 142-144.
Sullivan, Lucas, Heckert and Hunt, 1996. The type locality of Coelophysis,
a Late Triassic dinosaur from north-central New Mexico (USA). Palaeontologische
Zeitschrift. 70(1/2):245-255.
Sullivan and Lucas, 1999. Eucoelophysis baldwini, a new theropod dinosaur
from the Upper Triassic of New Mexico, and the status of the original types
of Coelophysis. Journal of Vertebrate Paleontology 19(1): 81-90.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur
record, part 3: Saurischians of North America. JVP. 25(3) 96A.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur
Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic
of New Mexico, USA. Geodiversitas. 28 (4),649-684.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
Coelophysidae indet. (Heckert, Zeigler, Lucas, Rinehart and Harris,
2000)
Norian, Late Triassic
Petrified Forest Formation of Chinle Group, New Mexico
Material- (NMMNH P-29168) tibia, fibula, astragalocalcaneum
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, 2732.
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, 127132.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
undescribed coelophysid (Taquet, 1984)
Toarcian, Early Jurassic
Morocco
Material- partial astragalus, calcaneum, distal tarsal III, metatarsal
I, phalanx I-1, ungual I, metatarsal II, phalanx II-1, phalanx II-2, ungual
II, metatarsal III (125 mm), phalanx III-1, proximal phalanx III-2, metatarsal
IV, phalanx IV-1, phalanx IV-2, metatarsal V
Comments- This is provisionally assigned to Coelophysidae based on the
apparent fusion between distal tarsal III and metatarsal III.
References- Jenny, Jenny-Deshusses, Le Marrec and Taquet, 1980. Decouverte
d'ossements de Dinosauriens dans le Jurassique inferieur (Toarcien) du Haut-Atlans
central (Maroc). Comptes Rendus Academie des Sciences. 290 839-842.
Taquet, 1984. Two new Jurassic specimens of coelurosaurs (Dinosauria). in Hecht,
Ostrom, Viohl and Wellnhofer (eds). The Beginnings of Birds: Proceedings of
the International Archaeopteryx Conference, Eichstaett. 229-232.
unnamed coelophysid (Fraser and Padian, 1995)
Norian?, Late Triassic
Pant-y-ffynnon fissure filling, Wales
Material- (BMNH PV RU 76/1, 77/1) posterior dorsal vertebrae, partial
sacrum, ilium (52 mm), incomplete pubis, partial ischium, incomplete femur
Comments- Rauhut and Hungerbuhler (2000) state it is extremely similar
to Megapnosaurus rhodesiensis in all features, although it is also very
similar to Procompsognathus and may prove to be referrable to that genus.
References- Fraser and Padian, 1995. Possible dinosaur remains from Britain
and the diagnosis of the Dinosauria. J.Vert.Paleont. 15(3): 30A.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia
15, 75-88.
Shake-N-Bake coelophysid (Tykoski, 1997)
Hettangian, Early Jurassic
Kayenta Formation, Arizona, US
Material- (MCZ 8817, 9442-9469; TMM 43689) (at least fifteen individuals,
adults) several hundred specimens including several dorsal vertebrae, several
sacra, many caudal vertebrae, several ilia, fifteen proximal femora, several
distal femora, six distal tibiotarsi (9.4, 11.1, 12.6, 12.9, 13 mm transversely),
including...
(MCZ 8817b) dorsal vertebra, partial sacrum, partial pelvis
(MCZ 8817c) two partial mid cervical vertebrae
(MCZ 8817d) incomplete posterior cervical vertebra
(MCZ 8817e) proximal caudal vertebra
(MCZ 8817f) partial tooth, partial caudal centrum
(MCZ 8817g) anterior dorsal vertebra
(MCZ 8817h) incomplete anterior dorsal vertebra
(MCZ 8817i) partial sacrum
(MCZ 8817j) proximal femur
(MCZ 8817k) (robust) proximal femur
(MCZ 8817l) (gracile) proximal femur
(MCZ 8817m) distal tibiotarsus
(MCZ 8817n) distal tibiotarsus, distal fibula
(MCZ 8817o) distal tibiotarsus, distal fibula
(MCZ 8817p) distal tibiotarsus
(MCZ 8817q) distal tibia, partial astragalus
(MCZ 8817r) partial scapulocoracoid
(MCZ 8817t) partial scapulocoracoid
(MCZ 9442; = 8817a) (adult) sacrum, partial ilia, proximal pubes, proximal ischia
(MCZ 9463; = 8817m-p?) (adult) distal tibiotarsus
(TMM 43689-4) (adult) proximal tarsometatarsus
Comments- This new taxon is based on specimens formerly thought to be
juvenile "Megapnosaurus" kayentakatae by Rowe (1989). Initially
catelogued at MCZ 8817 (Tykoski, 1998), at least some have since been recatalogued
as MCZ 9442-9469 and TMM 43689 (Tykoski, 2005). Tykoski (1998) used MCZ 8817a-s
as preliminary labels for some specimens, but it is unknown which numbers most
of these specimens are now catalogued under.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus
from the Early Jurassic Kayenta Formation of Arizona. J. Vertebr. Paleontol.
9 p. 125-136.
Tykoski, 1997. A new ceratosaurid theropod from the Early Jurassic Kayenta Formation
of Northern Arizona. JVP 17(3) 81A-82A
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications
for ceratosaurid phylogeny. Unpublished Masters Thesis,University of Texas at
Austin, 217 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
unnamed coelophysid (Lucas and Heckert, 2001)
Hettangian, Early Jurassic
Moenave Formation, Arizona, US
Material- (MNA V1968) several dorsal ribs or gastralia, incomplete ilium,
proximal pubis, proximal femur, incomplete ischium, distal ischium
(MNA V2588) five incomplete proximal caudal centra (34-37 mm), ilium (196 mm),
incomplete pubis, incomplete ischia (172 mm)
Comments- Pubes fuse to ischia posterior to acetabular midpoint, unlike
Coelophysis or Megapnosaurus species. Position ischia fuse to
ilia on antitrochanter more similar to "Megapnosaurus" kayentakatae
than Coelophysis. Medial bowing of femur more similar to M. rhodesiensis
than "M." kayentakatae.
References- Morales, 1994. First dinosaur body fossils from the Lower
Jurassic Dinosaur Canyon Member, Moenave Formation of Northeastern Arizona.
JVP. 14(3), 39A.
Lucas and Heckert, 2001. Theropod dinosaurs and the Early Jurassic age of the
Moenave Formation, Arizona-Utah, USA. Neues Jahrbuch fur Geologie und Palaontologie
Monatshefte 2001(7): 435-448.
unnamed coelophysid (Irmis, 2004)
Hettangian-Pliensbachian, Early Jurassic
Lower Lufeng Formation, Yunnan, China
Material- (FMNH CUP 2089) vertebra?, distal humerus, proximal radius,
proximal ulna, metacarpal fragments, manual phalangeal fragments
....(FMNH CUP 2090) distal tarsal II, distal tarsal III, proximal metatarsal
II, proximal metatarsal III
Comments- Simmons (1965) identified FMNH CUP 2089 as a podokesaurid distal
femur, proximal tibia and proximal fibula. Irmis (2004) referred it to Megapnosaurus
due to the fusion of metatarsals II and III, but the character is present in
Coelophysis bauri and the Shake-n-Bake coelophysid too (Tykoski, 2005).
References- Simmons, 1965. The non-therapsid reptiles of the Lufeng Basin,
Yunnan, China. Fieldiana: Geology 15:193.
Irmis, 2004. First report of Megapnosaurus (Theropoda: Coelophysoidea)
from China. PaleoBios. 24(3), 11-18.
undescribed Coelophysidae (UCMP online)
Late Carnian, Late Triassic
Petrified Forest Formation or Bluewater Creek Formation of the Chinle Group,
Arizona, US
Material- (UCMP 25962) ilium
(UCMP 25968) ilium
(UCMP 25971) pubis
(UCMP 25974) ilium
(UCMP 25978) ilium
(UCMP 25980) ischium
(UCMP 25982) ilium
(UCMP 25990) ischium
(UCMP 25993) ischium
(UCMP 25994) ischium
(UCMP 27982) ilium
Reference- U.C.M.P. Database. 2005. UCMP collections database. University
of California Museum of Paleontology.
undescribed Coelophysidae (UCMP online)
Late Triassic
Dockum Group, Texas, US
Material- (UCMP 65737) cervical vertebral fragments
(UCMP 65773) dorsal vertebra
(UCMP 65774) caudal vertebra
(UCMP 65775)
Reference- U.C.M.P. Database. 2005. UCMP collections database. University
of California Museum of Paleontology.
Coelophysis? longicollis
(Cope, 1887) Cope, 1889
= Coelurus longicollis Cope, 1887
= Tanystropheus longicollis Cope, 1887
Norian, Late Triassic
Petrified Forest Formation of the Chinle Group, New Mexico, US
Lectotype- (AMNH 2701) cervical vertebra (61 mm)
Comments- This was originally named based on the lectotype and four other
unassociated elements. 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 vertebra. 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 is probably indeterminate at the level
of Coelophysidae. The elongate centrum indicates it belongs to the clade.
References- Cope, 1887. The dinosaurian genus Coelurus. Amer.
Naturalist xxi pp. 367-369.
Cope, 1887. A contribution to the history of the Vertebrata of the Trias of
North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls. i, ii.
Cope, 1889. On a new genus of Triassic Dinosauria. Amer. Naturalist xxiii p.
626.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology
and comparisons. Palaeontographica 185A, 85-180.
Coelophysis? willistoni (Cope,
1887) Cope, 1889
= Tanystophaeus willistoni Cope, 1887
Norian, Late Triassic
Petrified Forest Formation of the Chinle Group, New Mexico, US
Lectotype- (AMNH 2726) partial ilium (160 mm)
Comments- This was originally named based on the lectotype and an unassociated
distal caudal centrum. Further material from the same locality was later included
in the hypodigm, but the lack of association prevents referring anything to
this taxon except the lectotype ilium. See the discussion of "unnamed Coelophysidae
(Cope, 1887)" above for more details. The lectotype is probably indeterminate
at the level of Coelophysidae. Although coelophysid synapomorphies aren't preserved
in the holotype, the coelophysoid-like angled pubic articular surface of the
ilium and association with coelophysid elements indicate the specimen was probably
a coelophysid itself.
References- Cope, 1887. A contribution to the history of the Vertebrata
of the Trias of North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls.
i, ii.
Cope, 1889. On a new genus of Triassic Dinosauria. Amer. Naturalist xxiii p.
626.
Longosaurus Welles, 1984
L. longicollis Welles 1984
Norian, Late Triassic
Petrified Forest Formation of the Chinle Group, New Mexico, US
Holotype- (AMNH 2705) ilium (189 mm)
Comments- Welles (1984) intended to separate Coelophysis longicollis
from Coelophysis bauri at the genus level by naming this genus, but accidentally
based it on a different specimen than the lectotype of Tanystropheus longicollis.
Thus these taxa have different type specimens and are not objective synonyms.
In addition, Welles referred at least some material Huene (1912) referred to
Coelophysis longicollis to Longosaurus longicollis (AMNH 2701,
2703, 2704, 2707). See the discussion of "unnamed Coelophysidae (Cope,
1887)" above for more details. The lectotype is probably indeterminate
at the level of Coelophysidae. The concave posterior postacetabular process
margin indicates it is a coelophysid however.
Reference- Cope, 1887. A contribution to the history of the Vertebrata
of the Trias of North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls.
i, ii.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology
and comparisons. Palaeontographica 185A, 85-180.
Podokesaurus Talbot, 1911
P. holyokensis Talbot, 1911
= Coelophysis holyokensis (Talbot, 1911)
Pliensbachian-Toarcian, Early Jurassic
Portland Formation?, Massachusetts, US
Holotype- (destroyed) (.89 m; .92 kg) cranial elements, four cervical
vertebrae (12-13 mm), cervical ribs, thirteen dorsal vertebrae (2nd 15 mm),
dorsal ribs, eleven rows of gastralia, thirteen caudal vertebrae (17 mm), coracoid,
humerus (42 mm), manus, ilial fragment, pubis (95 mm), ischium (50 mm), femur
(86 mm), tibia (104 mm), astragalus, metatarsal (65 mm), three partial metatarsals,
pedal digit (20 mm), gastrolith(?)
Pliensbachian-Toarcian, Early Jurassic
Portland Formation, Connecticut, US
Referred- ?(BSNH 13656) (2.5-3 m) tibiae, fibulae (Colbert and Baird,
1958)
Diagnosis- (after Colbert and Baird, 1958) dorsal neural spines anteroposteriorly
shorter than Coelophysis bauri.
Comments- Colbert and Baird (1958) found two differences from Coelophysis
bauri- dorsal neural spines anteroposteriorly shorter; ischium differently
shaped. The first is a good distinction (assuming it's real), but the second
certainly needs to be more precise. Podokesaurus may fall within the
genus Coelophysis or be indeterminate at the level of Coelophysidae,
but this is not formalized here pending redescription of C. bauri. It
is a coelophysid based on the elongate dorsal centra.
References- Talbot, 1911. Podokesaurus holyokensis, a new dinosaur
from the Triassic of the Connecticut Valley. Amer. Jour. Sci. 4 469-479, pl.
iv, 6 text-figs.
Colbert and Baird, 1958. Coelurosaur bone casts from the Connecticut Valley
Triassic. Am. Mus. Novitates 1901:1-11.
Pterospondylus Jaekel, 1913
P. trielbae Jaekel, 1913
Norian, Late Triassic
Trossingen Formation, Germany
Holotype- second dorsal vertebra (32 mm)
Comments- This species has been associated and synonymized with Procompsognathus
in the past based only on the elongate centrum. The transverse processes are
triangular, resembling Megapnosaurus more than Liliensternus and
Dilophosaurus. Thus, Rauhut and Hungerbuhler (2000) suggest the taxon
may be a coelophysid.
References- Jaekel, 1913.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia
15, 75-88.
Segisaurinae Camp, 1936 sensu Kalandadze
and Rautian, 1991
Diagnosis- (after Tykoski, 2005) anterodorsal margin of acromion has
smooth, continuous, high-angle transition to scapular blade; pubes rectangular
in distal view.
Comments- This subfamily was based on Camp's family Segisauridae and
is used here for the clade formed by Segisaurus, "Megapnosaurus"
kayentakatae and possibly Camposaurus recovered by Tykoski (2005).
Such a clade was not recovered by Ezcurra and Novas (2006), where "Megapnosaurus"
kayentakatae was instead closer to Coelophysis and Megapnosaurus
than to Segisaurus.
References- Camp, 1936. A new type of small bipedal dinosaur from the
Navajo sandstone of Arizona. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 24,
39-56.
Kalandadze and Rautian, 1991. Late Triassic zoogeography and reconstruction
of the terrestrial tetrapod fauna of North Africa. Paleont. Journ. 1, 1-12.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Camposaurus Hunt, Lucas,
Heckert, Sullivan and Lockley, 1998
C. arizonensis Hunt, Lucas, Heckert, Sullivan and Lockley, 1998
Late Carnian, Late Triassic
Mesa Redondo Formation of the Chinle Group, Arizona, US
Holotype- (UCMP 34498) distal tibiae, distal fibulae, astragalocalcanea
Referred- ?(UCMP 25791) pubic fragment, ilial fragment (Hunt,
Lucas, Heckert, Sullivan and Lockley, 1998)
Diagnosis- indeterminate relative to "Megapnosaurus" kayentakatae.
Comments- Originally referred to as Ceratosauria indet. by Long
and Murray (1995), UCMP 34498 was made the holotype of a new coelophysid taxon
by Hunt et al. (1998). Downs (2000) suggests that Camposaurus is a junior
synonym of Coelophysis, since it falls within the range of variation
of the Ghost Ranch specimens, and the initial comparisons were made with Colbert's
inaccurate monograph. Nesbitt et al. (2005, 2007) agree it cannot be distinguished
from Coelophysis bauri, as the straight ventral astragalar margin is
found in the latter as well.. Tykoski (2005) notes the presence of fibulocalcanear
fusion is otherwise only seen in "Megapnosaurus" kayentakatae,
though otherwise the material cannot be placed more precisely within the Coelophysis<Liliensternus
clade.
UCMP 25791 was figured by Long and Murray, then assigned questionably to Camposaurus
by Hunt et al.. Tykoski noted the pelvic fragment indicates the ilium was fused
to the pubis, which is a ceratosaurian character. Thus it may be properly referred
to Camposaurus.
A proximal femur (UCMP 139662), dorsal centra (MNA V3091; UCMP 177317), and
sacral remains (UCMP 138591, 178047-178049) were identified as Ceratosauria
indet. by Long and Murray, then made paratypes of Camposaurus
by Hunt et al.. The latter authors felt the material may even belong to the
holotype individual, but Nesbitt et al. ciuld only identify the femur to Saurischia
indet. and the axial material to Archosauria indet..
References- Long and Murry, 1995. Late Triassic (Carnian and Norian)
tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History
Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from
the western United States. Geobios, 31(4):511-531.
Downs, 2000. Coelophysis bauri and Syntarsus rhodesiensis compared,
with comments on the perparation and preservation of fossils from the Ghost
Ranch Coelophysis quarry. in Lucas and Heckert (eds.), 2000. Dinosaurs
of New Mexico. NMMNH Bulletin 17. 33-37.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur
record, part 3: Saurischians of North America. JVP. 25(3) 96A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209243.
"Megapnosaurus" kayentakatae
(Rowe, 1989) Tykoski and Rowe, 2004
= Syntarsus kayentakatae Rowe, 1989
= Coelophysis kayentakatae (Rowe, 1989) Bristowe and Rowe, 2004
Hettangian, Early Jurassic
Silty Facies Member of the Kayenta Formation, Arizona, US
Holotype- (MNA V2623) (robust adult) skull (230 mm), mandibles, hyoids,
atlas, axis (27.8 mm), third cervical vertebra (33 mm), fourth cervical vertebra
(>41 mm), fifth cervical vertebra, sixth cervical vertebra (>45 mm), seventh
cervical vertebra (46.7 mm), eighth cervical vertebra (43.5 mm), ninth cervical
vertebra, tenth cervical vertebra (>33 mm), cervical ribs 1-10 (110-295 mm),
first dorsal vertebra, partial second dorsal vertebra, first dorsal rib, over
fifteen rows of gastralia, two partial sacral centra, caudal vertebrae, scapulocoracoids
(one partial; 181 mm), furcula, incomplete humerus (~116 mm), distal carpal
I+II, proximal carpal, carpal, metacarpal I (21.2 mm), manual ungual I (21 mm),
phalanx II-2 (24 mm), proximal manual ungual II, metacarpal III (>39 mm),
phalanx III-3 (18.8 mm), manual ungual III (18.2 mm), metacarpal IV (>20.8
mm), phalanx IV-1 (13 mm), phalanx IV-2, partial pubes, partial ischia, femora
(one partial) (276 mm), tibiae (one partial) (292 mm), fibulae (one partial)
(274 mm), astragalocalcaneum (32.2, 33.5 mm transversely), distal tarsal III,
distal tarsal IV, metatarsal I (38 mm), phalanx I-1 (27.2, 27.3 mm), pedal ungual
I (15, 16 mm), metatarsal II (150 mm), phalanx II-1 (47, 44.8 mm), phalanx II-2
(38 mm), pedal ungual II (30 mm), metatarsal III (173 mm), phalanx III-1 (50,
47.7 mm), phalanx III-2 (33, 36.9 mm), phalanx III-3 (34, 34.6 mm), pedal ungual
III (28.5 mm), metatarsal IV, phalanx IV-1 (27.5 mm), phalanx IV-2, phalanx
IV-3 (18.3, 18.4 mm), phalanx IV-4 (15.5, 15.6 mm), pedal ungual IV (>24
mm), metatarsal V (>57 mm)
Paratypes- (MNA V2623) (15-20% larger than holotype; at least two robust)
snout, dentaries, maxillary fragment with ninth to eleventh teeth, frontals,
partial occipital condyle, fragments of sacra, portions of ~34 caudal centra,
two fragmentary femora, three proximal tibiae, three astragalocalcanea, eight
ends of metatarsals
(MCZ 8817) (at least eleven robust individuals) caudal, pelvic and hindlimb
elements
?(MNA V100) partial ilium
?(MNA V140) (juvenile) humerus, femur, partial tibia, bone fragments
(TMM 43648-9)
(TMM 43669-3) (young subadult)
(TMM 43688-1) (robust old subadult) cervical vertebrae, sacrum (27, 23.4, 22.2,
24, 26 mm = 121 mm), few caudal centra, partial ilia, pubes (<232 mm), ischia
(121 mm), femora (229.8, 230 mm), proximal tibia, proximal fibula, distal tarsal
III, distal tarsal IV, proximal tarsometatarsus, several pedal phalanges
(UCMP V128659) (gracile) several proximal caudal centra, fragmentary ilium,
proximal pubes, proximal femora, proximal fibula
Diagnosis- (after Tykoski and Rowe, 2004) nasal crests; frontals separated
by midline anterior extension of parietals.
Comments- The 'juveniles' previously referred (Rowe, 1989) are actually
the distinct "Shake-n-Bake" coelophysid (Tykoski, 1998). The sacral
and caudal material assigned to the holotype may belong to the other two individuals
with the same specimen number. Tykoski 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). Tykoski (2005) found this species to be more closely related
to Segisaurus than to Coelophysis or Megapnosaurus.
References- Rowe, 1989. A new species of the theropod dinosaur Syntarsus
from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate
Paleontology. 9, 125-136.
Tykoski, 1998. The osteology of Syntarsus kayentakatae and its implications
for ceratosaurid phylogeny. Unpublished Masters Thesis, University of Texas
at Austin, 217 pp.
Ivie, Slipinski and Wegrzynowicz, 2001. Generic homonyms in the Colydiinae (Coleoptera:
Zopheridae). Insecta Mundi 15:63-64.
Tykoski, Forster, Rowe, Sampson and Munyikwa, 2002. A furcula in the coelophysid
theropod Syntarsus. Journal of Vertebrate Paleontology. 22(3):728733.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The
Dinosauria Second Edition. University of California Press. 861 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Segisaurus Camp, 1936
S. halli Camp, 1936
Toarcian, Early Jurassic
Navajo Sandstone, Arizona, US
Holotype- (UCMP 32101) (robust old subadult) (1.54 m) distal cervical
ribs 7-10, partial fourth dorsal vertebra, partial fifth dorsal vertebra, partial
sixth dorsal vertebra, partial eighth dorsal vertebra, partial ninth dorsal
vertebra, dorsal neural arch, several partial dorsal ribs, over twelve rows
of gastralia, impression of first sacral centrum, impression of second sacral
centrum, partial third sacral vertebra, partial fouth sacral vertebra, partial
caudal vertebrae 2-22, fragments of several caudal vertebrae, several chevrons,
scapula (93 mm), partial coracoids, partial furcula, possible sternal fragment,
incomplete humerus (72 mm), proximal radius, proximal ulna, distal phalanx II-2,
manual ungual II, distal phalanx III-2, phalanx III-3, manual ungual III, manual
phalanges, ilial fragments, incomplete pubes (~110 mm), incomplete ischia (~96
mm), incomplete femora (~145 mm), tibiae (160 mm), incomplete fibula, fibular
fragment, partial astragalus, calcaneum, distal tarsal IV, metatarsal I (33
mm), phalanx I-1 (17 mm), pedal ungual I, metatarsals II, phalanx II-1, phalanx
II-2 (23 mm), pedal ungual II, metatarsals III (99 mm), phalanx III-1, phalanx
III-2 (22 mm), phalanx III-3 (13 mm), pedal ungual III (10 mm), metatarsal IV,
phalanx IV-1, phalanx IV-2 (14 mm), phalanx IV-3, phalanx IV-4, pedal ungual
IV (14 mm), metatarsals V (32 mm)
Diagnosis- (from Rauhut, 2003) dorsal centra not very constricted ventrally;
slender scapula; humeral shaft with stronger torsion than Coelophysis
(~50 degrees); large ischial fenestra; (from Carrano et al., 2005) rectangular
humeral deltopectoral crest.
Comments- Segisaurus has been reprepared, showing a furcula in
articulation with the coracoids (not separate clavicles) and thin-walled long
bones.
Although Carrano et al. (2005) could not recover any resolution within Coelophysoidea
in their cladistic analysis, when several characters are changed to ordered,
the results differ. If a standard coelophysoid topology of (Dilophosaurus
(Liliensternus (Coelophysis, Megapnosaurus))) is enforced,
Segisaurus is placed outside the Liliensternus + Coelophysis
+ Megapnosaurus clade. This differs from Rauhut's (2000) findings, which
placed it in the Coelophysidae.
References- Camp and VanderHoof, 1935. Small bipedal dinosaur from the
Jurassic of northern Arizona. Proc. Geol. Soc. Amer. 1934 384-385.
Camp, 1936. A new type of small bipedal dinosaur from the Navajo sandstone of
Arizona. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 24, 39-56.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.], 1-440.
Senter and Hutchinson, 2001. New information on the skeleton of the theropod
Segisaurus halli. JVP 21(3) 100A.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli,
a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate
Paleontology. 25(4), 835849.
Coelophysinae Nopcsa, 1928
Definition- (Coelophysis bauri <- Segisaurus halli)
(proposed)
Other definitions- (Coelophysis bauri <- Procompsognathus
triassicus) (Sereno, in press; modified from Sereno, 1998)
= Syntarsiinae Kalandadze and Rautian, 1991
Diagnosis- (modified from Tykoski, 2005) dorsal process of maxilla angles
<35 degrees from horizontal; promaxillary fenestra absent; lateral lamina
of ventral lacrimal process linear and remains posterior to medial lamina; more
than 18 dentary teeth; jugal does not contact antorbital fossa.
Comments- Tykoski (2005) also included the reduction in size of dorsolateral
nasal ridges as a synapomorphy, but as Zupaysaurus' ridges are more likely
to be displaced nasals, this becomes an ambiguous character not favored by DELTRAN.
It is viewed as evolving convergently in Dilophosaurus and "Megapnosaurus"
kayentakatae here.
Alhough originally proposed by Nopcsa in 1928, this subfamily went largely unused
until Paul (1988) used it for Coelophysis, Megapnosaurus rhodesiensis
and Elaphrosaurus, to separate them from the halticosaurine Liliensternus
and Dilophosaurus. Sereno (1998) then used it to group Coelophysis
and Megapnosaurus together, defining it to include Coelophysis
and exclude Procompsognathus. In Sereno's topology, this also excluded
Segisaurus and Liliensternus. Yet subsequent studies (e.g. Tykoski,
2005; Ezcurra and Rautian, 2006) have shown Procompsognathus to have
an uncertain placement within derived coelophysoids, making the precise application
of Sereno's definition impossible and the content of his Coelophysinae (besides
Coelophysis itself) unknown. Carrano et al. (2002) later used Coelophysinae
for a Megapnosaurus + Coelophysis clade, excluding Liliensternus.
Taking these prior uses into account, Coelophysinae is here used for the Megapnosaurus
+ Coelophysis clade, but with Segisaurus as the external specifier
instead of Procompsognathus. This is a clade recovered in most phylogenetic
analyses where coelophysid interrelationships are resolved (e.g. Tykoski, 2005;
Ezcurra and Novas, 2006) and is equivalent to Sereno's Coelophysinae
in content.
Kalandadze and Rautian (1991) proposed the subfamily Syntarsiinae, but this
cannot be used for a theropod group, as Syntarsus is the name of a beetle
(Ivie et al., 2001).
References- Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1,
163-188.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Kalandadze and Rautian, 1991. Late Triassic zoogeography and reconstruction
of the terrestrial tetrapod fauna of North Africa. Paleont. Journ. 1, 1-12.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und
Paläontologie, Abhandlungen. 210(1), 41-83.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri,
a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar.
Journal of Vertebrate Paleontology. 22(3), 510-534.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Ezcurra and Novas, 2006. Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NWArgentina. Historical Biology. iFirst Article,
38 pp. DOI 10.1080/08912960600845791.
Coelophysidae sensu Tykoski and Rowe, 2004
Definition- (Coelophysis bauri + Megapnosaurus rhodesiensis) (modified)
Coelophysinae sensu Sereno, 1998
Definition- (Coelophysis bauri <- Procompsognathus triassicus)
Coelophysis Cope, 1889
= Rioarribasaurus Hunt and Lucas, 1991
C. bauri (Cope, 1887) Cope, 1889
= Coelurus bauri Cope, 1887
= Tanystropheus bauri (Cope, 1887) Cope, 1887
= Rioarribasaurus colberti Hunt and Lucas, 1991
= Syntarsus colberti (Hunt and Lucas, 1991) Paul, 1993
Norian, Late Triassic
Rock Point Formation of the Chinle Group, 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),
femur (203 mm), tibia (221 mm), astragalus, calcaneum, (astragalocalcaneum 23
mm wide), metatarsus (120 mm)
Referred- (AMNH 7223) (2.68 m, 15.3 kg) complete skeleton including skull
(265 mm), cervical series (485 mm), dorsal series (425 mm), humerus (120 mm),
femur (209 mm), tibia (224 mm), astragalus, calcaneum, metatarsal II (110 mm),
metatarsal III (126 mm), metatarsal IV (114 mm)
(AMNH 7225) skull (Colbert, 1989)
(AMNH 7226) pes (Colbert, 1989)
(AMNH 7227) skull (159 mm), cervical series (384 mm), dorsal series (360 mm),
sacrum, scapulae, humeri (86 mm), radii, ulnae, manus, ilium, pubis, proximal
ischium, femora (165 mm), tibiae, fibulae, pes (Colbert, 1989)
(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,
1989)
(AMNH 7229) six dorsal vertebrae, sacral vertebrae, caudal vertebrae, partial
ilia, pubis, ischium, femora (135 mm), tibiae (154 mm), fibulae, metatarsus
(85 mm), pedal phalanges (Colbert, 1989)
(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,
femora (125 mm), tibiae, fibulae (Colbert, 1989)
(AMNH 7231) incomplete cervical series (215 mm), incomplete dorsal series (262
mm), sacrum, scapulae, coracoids, radius, ulna, partial ilium, pubis, femora
(132 mm), tibiae, fibulae, pes (Colbert, 1989)
(AMNH 7232) ilium, pubes, femur (141 mm), tibia (157 mm), fibula, metatarsus
(95 mm), pedal phalanges (Colbert, 1989)
(AMNH 7233) dorsal vertebrae, sacral vertebrae, ilium, pubis, ischium, femora
(126 mm), tibiae (140 mm), fibulae, metatarsus (81 mm), pedal phalanges (Colbert,
1989)
(AMNH 7234) dorsal vertebrae, sacral vertebrae, partial pubis, ischium, tibia
(135 mm), fibula, metatarsus (84 mm), pedal phalanges (Colbert, 1989)
(AMNH 7235) ilium, ischium, partial femur (Colbert, 1989)
(AMNH 7236) ilium, pubis, femur (Colbert, 1989)
(AMNH 7237) skull, cervical vertebrae (Colbert, 1989)
(AMNH 7238) skull (Colbert, 1989)
(AMNH 7239) skull, mandibles, cervical vertebrae (Colbert, 1989)
(AMNH 7240) skull (198 mm), mandibles (Colbert, 1989)
(AMNH 7241) skull (140 mm), mandibles (Colbert, 1989)
(AMNH 7242) (juvenile) skull (~68 mm) (Colbert, 1989)
(AMNH 7243) ten dorsal vertebrae, dorsal ribs, sacrum, proximal caudal vertebrae,
scapulacoracoid, humeri (91 mm), radii, ulnae, manus, pelvis, femora (172 mm),
tibiae (199 mm), fibulae, astragalus, calcaneum, metatarsus (114 mm), pedal
phalanges (Colbert, 1989)
(AMNH 7244) proximal thirty caudal vertebrae, pelvis, femora (173 mm), tibiae
(195 mm), fibulae, metatarsus (120 mm), pedal phalanges (Colbert, 1989)
(AMNH 7245) five or six presacral vertebrae, six proximal caudal vertebrae,
pelvis, femur, tibia (Colbert, 1989)
(AMNH 7246) eight caudal vertebrae, pelvis, femur (122 mm), tibiae (136 mm),
fibulae, metatarsi (79 mm), pedal phalanges (Colbert, 1989)
(AMNH 7247) femur (125 mm), tibia (138 mm), metatarsus (84 mm), pedal phalanges
(Colbert, 1989)
(AMNH 7248) ilium, pubis (Colbert, 1989)
(AMNH 7249) eight dorsal vertebrae, dorsal ribs, sacrum, five caudal vertebrae,
pelvis, femora (196 mm), tibiae (207 mm), fibulae, metatarsi (110 mm), pedal
phalanges (Colbert, 1989)
(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,
1989)
(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,
1989)
(AMNH 7257) seventeen presacral vertebrae, humeri, partial radii, partial ulnae
(Colbert, 1989)
(AMNH 7258) skull, mandible, cervical vertebrae 1-7, seven or eight dorsal vertebrae
(Colbert, 1989)
(CM 31374) skull, mandible (Downs, 2000)
(CM C-1-82) (one or two individuals) partial skeleton (Colbert, 1989)
(CM C-3-82) (juvenile) skeleton including nasal, vertebrae, furcula, pelvis,
hindlimbs (Downs, 2000)
(CM C-4-81) (at least nine individuals) skull (250 mm), maxillary fragment,
caudal vertebrae 1-5, two pelves, two hindlimbs, femur, distal tibiae, two pes
(Colbert, 1989)
(GR141) (Downs, 2000)
(GR142) (Downs, 2000)
(GR1442) (Downs, 2000)
(MCZ 4326) skull, mandible (Colbert, 1989)
(MCZ 4327) skull (239 mm), mandible, atlas, axis, pubis, femur, proximal tibia
(Colbert, 1989)
(MCZ 4328) partial maxilla (Colbert, 1989)
(MCZ 4329) forelimb including five carpals (Colbert, 1989)
(MCZ 4330) pelvis (Colbert, 1989)
(MCZ 4331) (first individual) last thirteen dorsal vertebrae, dorsal ribs, humerus,
radius, ulna, pelvis, hindlimbs
(second individual) two distal caudal vertebrae, femur, tibia, other elements
(third individual) three cervical vertebrae, three dorsal vertebrae, pelvis,
femora, tibia (Colbert, 1989)
(MCZ 4332) (three or four individuals) vertebrae, pelves, hindlimbs (Colbert,
1989)
(MCZ 4333) skull, partial mandible, anterior cervical vertebrae, partial manus
(Colbert, 1989)
(MCZ 4334) vertebrae, pelvis, hindlimbs (Colbert, 1989)
(MCZ 4335) vertebrae (Colbert, 1989)
(MNA V3139) fragmentary skull, fragmentary mandibles, postcrania (Colbert, 1989)
(MNA V3315) skull (143 mm), mandibles, atlas, axis, third cervical vertebra
(Colbert, 1989)
(MNA V3318) (1.8 m) incomplete skeleton including skull (~88 mm), cervical series
(184 mm), dorsal series (255 mm), femur (123 mm), tibia (136 mm), astragalocalcaneum,
metatarsal II (72 mm), metatarsal III (82 mm), metatarsal IV (70.5 mm) (Colbert,
1989)
(MNA V3319) pes including metatarsal II (67 mm), metatarsal III (76 mm), metatarsal
IV (69 mm) (Colbert, 1989)
(MNA V3320) pes including astragalus, calcaneum, metatarsal II (115 mm), metatarsal
III (128 mm), metatarsal IV (118 mm) (Colbert, 1989)
(MNA V3321) distal tibia, astragalocalcaneum (Colbert, 1989)
(MNA V3322) partial skull (Colbert, 1989)
(MNA V3323) caudal series (Colbert, 1989)
(NMMNH P-42352) skeleton including cololite and coprolite (Rinehart, Hunt, Lucas,
Heckert and Smith, 2005)
(NMMNH P-44552) skeleton including coprolite with juvenile elements included
(rib fragments, ulnare, proximal metacarpals, partial phalanges, long bone fragments,
cranial and/or pelvic material) (Rinehart, Hunt, Lucas, Heckert and Smith, 2005)
(NMMNH P-44801) skeleton including coprolite (Rinehart, Hunt, Lucas, Heckert
and Smith, 2005)
(NMMNH P-C-8-82) (several individuals including three juveniles) material including
five furculae (Rinehart, Lucas and Hunt, 2006)
(RTMP 84-63-23) skull (Colbert, 1989)
(RTMP 84-63-29) skull, mandibles, cervical vertebrae, dorsal vertebrae (Colbert,
1989)
(RTMP 84-63-30) cervical vertebrae, dorsal vertebrae, scapula, coracoid, humerus,
radius, ulna, manus (Colbert, 1989)
(RTMP 84-63-31) maxilla, mandible (Colbert, 1989)
(RTMP 84-63-32) anterior skull, anterior mandible, posterior cervical vertebrae,
dorsal vertebrae, ribs, scapulocoracoid, forelimbs (Colbert, 1989)
(RTMP 84-63-33) last thirteen dorsal vertebrae, ribs, sacrum, more than fifteen
caudal vertebrae, scapula, coracoid, humerus, pelvis, hindlimbs including metatarsal
II (94.2 mm), metatarsal III (105 mm), metatarsal IV (93 mm) (Colbert, 1989)
(RTMP 84-63-34) last seven dorsal vertebrae, several caudal vertebrae, pelvis,
femora, tibiae, fibula, astragalocalcaneum, pes (Colbert, 1989)
(RTMP 84-63-35) eleven caudal vertebrae, humerus(?), tibia, fibula, astragalus,
calcaneum, pes (Colbert, 1989)
(RTMP 84-63-36) vertebrae, femora, tibiae, other elements (Colbert, 1989)
(RTMP 84-63-37) vertebrae, scapulacoracoid(?), ischia, hindlimbs (Colbert, 1989)
(RTMP 84-63-38) pelvis (Colbert, 1989)
(RTMP 84-63-39) vertebrae, ilium, femur (Colbert, 1989)
(RTMP 84-63-40) manus (Colbert, 1989)
(RTMP 84-63-41) six caudal vertebrae (Colbert, 1989)
(RTMP 84-63-42) partial femora, tibiae (Colbert, 1989)
(RTMP 84-63-43) seven series of caudal vertebrae (Colbert, 1989)
(RTMP 84-63-44) vertebrae, other elements (Colbert, 1989)
(RTMP 84-63-45) fragmentary skull (Colbert, 1989)
(RTMP 84-63-46) manus (Colbert, 1989)
(RTMP 84-63-47) distal tibia, distal fibula, astragalocalcaneum (Colbert, 1989)
(RTMP 84-63-48) sacrum, posterior ilia (Colbert, 1989)
(RTMP 84-63-49) eleven vertebrae (Colbert, 1989)
(RTMP 84-63-50) manus (Colbert, 1989)
(RTMP 84-63-51) dorsal vertebrae, dorsal ribs (Colbert, 1989)
(RTMP 84-63-52) carpus, manus (Colbert, 1989)
(RTMP 84-63-53) seven cervical vertebrae (Colbert, 1989)
(RTMP 84-63-54) ten caudal vertebrae (Colbert, 1989)
(RTMP 84-63-55) femur (Colbert, 1989)
(RTMP 84-63-56) fourth distal tarsal, metatarsal (Colbert, 1989)
(RTMP 84-63-57) vertebrae (Colbert, 1989)
(RTMP 84-63-58) caudal vertebrae, long bone (Colbert, 1989)
(RTMP 84-63-59) six posterior cervical vertebrae (Colbert, 1989)
(RTMP 84-63-60) element (Colbert, 1989)
(RTMP 84-63-61) vertebrae, two metatarsals (Colbert, 1989)
(YPM 41196) skull (211 mm), mandible (Colbert, 1989)
(YPM 41197) pelvis (Colbert, 1989)
(YPM 41412) pelvis, hindlimb (Colbert, 1989)
(YPM 43506) anterior skull (Colbert, 1989)
Norian, Late Triassic
Petrified Forest Formation of the Chinle Group, Arizona, US
(PEFO 33981) fragmentary skeleton including partial posterior vertebral column,
pelvis, incomplete hindlimbs (Nesbitt et al., 2007)
(PEFO 33983) incomplete posterior skeleton (Nesbitt et al., 2007)
(UCMP 129618) (3.46 m) dorsal centrum (45 mm), partial dorsal centrum, three
partial sacral centra, proximal caudal centrum (33 mm), partial ilium, nearly
complete pubis, partial ischium, femora (235, 245 mm), tibia (255 mm), fibula,
astragalocalcaneum, distal tarsal III, distal tarsal IV, phalanx I-1 (29 mm),
metatarsal III (150 mm), proximal phalanx III-1, phalanx III-2 (37 mm), distal
pedal ungual III, metatarsal IV (133 mm), phalanx IV-1 (30 mm), phalanx IV-3
(22 mm), phalanx IV-4 (19 mm), proximal pedal ungual IV (Padian, 1986)
partial skeleton including teeth, cervical vertebrae, femora and tibiae (Hunt
and Wright, 1999)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, Texas, US
?(TTUP 10071) partial ilium (Lehman and Chatterjee, 2005)
?(TTUP 10072) vertebrae, proximal femur, distal tibia, astragalus (Lehman and
Chatterjee, 2005)
?(TTUP 10082) pubis (Lehman and Chatterjee, 2005)
Late Triassic(?)
US(?)
(CM 11894) tibia, fibula, astragalus, calcaneum (Tykoski, 2005)
(SMP VP-453) (Sullivan et al., 1996)
(SMP VP-456) (Sullivan et al., 1996)
(SMP VP-462) (Sullivan et al., 1996)
(SMP VP-465) (Sullivan et al., 1996)
(SMP VP-469) (three individuals) (Sullivan et al., 1996)
(SMP VP-476) (Sullivan et al., 1996)
(SMP VP-478) (four specimens) (Sullivan et al., 1996)
(SMP VP-487) (Sullivan et al., 1996)
(SMP VP-491) (Sullivan et al., 1996)
(SMP VP-1072) femur
(TMM 45559) cervical vertebrae, sacral vertebrae, partial ilium, femur, tibia,
astragalocalcaneum (Tykoski, 2005)
Removed material- Several authors (Gregory, 1945; Elder, 1978, 1987)
assigned material from the Colorado City Member of the Dockum Formation in Texas
to Coelophysis, but these are Trilophosaurus (Hunt et al., 1998).
The vertebral centra from the Blue Mesa Formation of the Chinle Group referred
to Coelophysis by Colbert (1989) are Trilophosaurus (Hunt et al.,
1998).
Diagnosis- (after Downs, 2000) unspecified differences from Megapnosaurus
in cervical length, proximal and distal hindlimb proportions and proximal
caudal vertebral anatomy.
(after Tykoski and Rowe, 2004) differs from Megapnosaurus and "M."
kayentakatae in lacking pit at the base of the nasal process of the premaxilla.
(after Bristowe and Raath, 2004) differs from Megapnosaurus in having
a longer maxillary tooth row; anteroposterior length of ventral lacrimal process
>30% of its height.
(after Ezcurra, 2006) absence of an offset rostral process of the maxilla; strongly
caudally bowed quadrate; small external mandibular fenestra (9-10% of mandibular
length) (also in Dilophosaurus). Differs from Megapnosaurus in
having a square-shaped rostral process of the jugal; unreduced medial condyle
of the distal trochlea of the metacarpal I.
Comments- Hunt and Lucas (1991) felt Cope's original Coelophysis
specimens (including the lectotypes of C. bauri, C. longicollis
and C. willistoni) were indeterminate, and created the name Rioarribasaurus
colberti for the abundant complete Ghost Ranch specimens. However, the ICZN
later (1996) ruled a Ghost Ranch specimen is the neotype of Coelophysis bauri.
Coelophysis? longicollis, C? willistoni, the other original Coelophysis
specimens, Longosaurus, Podokesaurus and Protoavis (in
part) may all be referrable to C. bauri or at least Coelophysis,
but this cannot be determined until the Ghost Ranch specimens are redescribed.
The illustrations of the latter in Colbert (1989) are inaccurate and have hampered
comparison to other coelophysids. Downs (2000) notes most of the supposed differences
from Megapnosaurus rhodesiensis are not real (C. bauri has a nasal
fenestra; vaulted palate; interdental plates; obturator fenestra; pubic fenestra;
fused pelvis; triangular dorsal transverse processes; identical hindlimb morphology).
Tykoski (2005) notes the presence of a median basisphenoid spur that follows
the roof of the basisphenoid recess cannot be ascertained in Coelophysis
(contra Tykoski and Rowe, 2004). Ezcurra (2006) noted that contra Bristowe and
Raath (2004), an anteriorly pointed antorbital fossa only occurs in some adult
specimens (e.g. CM C-3-82, AMNH 7224, YPM 41196), but not others (e.g. AMNH
7240, MCZ 4327). It is thus not a diagnostic character of the species.
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.
References- Cope, 1887. The dinosaurian genus Coelurus. Amer.
Naturalist xxi pp. 367-369.
Cope, 1887. A contribution to the history of the Vertebrata of the Trias of
North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls. i, ii.
Cope, 1889. On a new genus of Triassic Dinosauria. Amer. Naturalist xxiii p.
626.
Huene, 1915. On reptiles of the New Mexican Trias in the Cope collection. Bulletin
American Museum of Natural History. 34, 485-507.
Colbert and Baird, 1958. Coelurosaur bone casts from the Connecticut Valley
Triassic. Amer. Mus. Novit. no. 1901 11 pp., 2 figs., 1 table.
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). p. 45-60 in Kevin Padian (ed.), The beginning of the Age of Dinosaurs.
Faunal change across the Triassic-Jurassic boundary. Cambridge University Press,
Cambridge, England.
Colbert, 1989. The Triassic dinosaur Coelophysis. Museum of Northern
Arizona Bulletin. 57:1-174.
Colbert, 1990. Variation in Coelophysis bauri. p. 81-90 in Kenneth Carpenter,
Philip J. Currie (eds.), Dinosaur systematics. Approaches and perspectives.
Cambridge University Press, Cambridge, etc..
Rowe and Gauthier, 1990. Ceratosauria. p. 151-168. in David B. Weishampel, et
al. (eds.), The Dinosauria. University of California Press, Berkeley, Los Angeles,
Oxford.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic
dinosaur from New Mexico (USA). Paläontol. Z. 65 p. 191-198.
Colbert, Harris, Charig, Dodson, Gillette, Ostrom and Weishampel, 1992. Coelurus
bauri Cope, 1887 (currently Coelophysis bauri; Reptilia, Saurischia):
Proposed replacement of the lectotype by a neotype. Bull. Zool. Nomencl. 49
p. 276-279.
International Commision on Zoological Nomenclature, 1996. Opinion 1842. Coelurus
bauri Cope, 1887 (currently Coelophysis bauri; Reptilia, Saurischia):
lectotype replaced by a neotype. Bulletin of Zoological Nomenclature. 53, 142-144.
Sullivan, Lucas, Heckert and Hunt, 1996. The type locality of Coelophysis,
a Late Triassic dinosaur from north-central New Mexico (USA). Palaeontologische
Zeitschrift. 70(1/2):245-255.
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 McClellan, eds.. National Park
Service Paleontological Research. National Park Service Geological Resources
Division Technical Report. NPS/NRGRD/GRDTR-99/03, p. 96-99.
Downs, 2000. Coelophysis bauri and Syntarsus rhodesiensis compared,
with comments on the perparation and preservation of fossils from the Ghost
Ranch Coelophysis quarry. in Lucas and Heckert (eds.), 2000. Dinosaurs
of New Mexico. NMMNH Bulletin 17. 33-37.
Smith, Andersen, Larsson and Bybee, 2000. Results of a high resolution CT-scan
of Coelophysis bauri (Upper Triassic Chinle Formation, New Mexico). JVP.
20(3) 70A.
Rinehart, Lucas and Heckert, 2001. Preliminary statistical analysis defining
the juvenile, robust and gracile forms of the Triassic dinosaur Coelophysis.
JVP. 21(3) 93A.
Gay, 2002. The myth of cannibalism in Coelophysis bauri. J. Vert. Paleontol.
22, 57A.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska. The
Dinosauria Second Edition. University of California Press. 861 pp.
Lehman and Chatterjee, 2005. Depositional setting and vertebrate biostratigraphy
of the Triassic Dockum Group of Texas. J. Earth Syst. Sci.. 114(3), 325351.
Parker and Irmis, 2005. Advances in Late Triassic vertebrate paleontology based
on new material from Petrified Forest National Park, Arizona. Vertebrate Paleontology
in Arizona, New Mexico Museum of Natural History and Science Bulletin. 29:45-58.
Rinehart, Hunt, Lucas and Heckert, 2005. Coprolites and cololites from the Late
Triassic theropod dinosaur Coelophysis bauri, Whitaker Quarry, Rio Arriba
County, NM. New Mexico Geology. 27(2), 53.
Rinehart, Hunt, Lucas, Heckert and Smith, 2005. New evidence of cannibalism
in the Late Triassic (Apachean) dinosaur, Coelophysis bauri (Theropoda:
Ceratosauria). JVP. 25(3) 105A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. The cranial anatomy of the coelophysoid theropod Zupaysaurus
rougieri from the Upper Triassic of Argentina. Historical Biology. iFirst
Article, 18 pp. DOI 10.1080/08912960600861467.
Nesbitt, Turner, Erickson and Norell, 2006. Prey choice and cannibalistic behaviour
in the theropod Coelophysis. Biol. Lett., 4 pages. doi:10.1098/rsbl.2006.0524.
Published online.
Rinehart, Lucas and Hunt, 2006. The furcula of Coelophysis bauri, a Late
Triassic (Apachean) dinosaur (Theropoda: Ceratosauria) from New Mexico. New
Mexico Geology. 28(2), 62.
Megapnosaurus Ivie, Slipinski and Wegrzynowicz, 2001
= Syntarsus Raath, 1969 (preoccupied Fairmaire, 1869)
Comments- This genus is often synonymized with Coelophysis (Paul,
1988; Bristowe and Raath, 2004), as new discoveries invalidate supposed differences
between the taxa (e.g. Downs, 2000; Bristowe and Raath, 2004). In 2001, it was
discovered that Syntarsus was preoccupied by a beetle. The entomologists
who determined this attempted unsuccessfully to contact Raath so that he could
rename it, and ended up renaming it themselves (Ivie et al., 2001). Paleontologists
might have reacted more positively if the replacement name (Megapnosaurus)
hadn't been facetious, translating to "big dead lizard", since that's
what all dinosaurs are to entomologists. Thus the trend has been to ignore the
name, a position made easier by its resemblence to Coelophysis. However,
personal dislike for a name is not a valid reason to reject it, and Coelophysis
and Megapnosaurus are more closely related to each other than either
is to other well known coelophysoids (Procompsognathus, Segisaurus,
Liliensternus, Gojirasaurus, etc.). Thus their synonymy is a subjective
decision, which some workers follow (Downs, 2000; Bristowe and Raath, 2004)
and others do not (Irimis, 2004; Tykoski and Rowe, 2004). I keep them separate
primarily due to the influence of Tykoski and Rowe's paper.
M. rhodesiensis (Raath,
1969) Ivie, Slipinski and Wegrzynowicz, 2001
= Syntarsus rhodesiensis Raath, 1969
= Coelophysis rhodesiensis (Raath, 1969) Paul, 1988
Hettangian-Sinemurian, Early Jurassic
Upper Elliot Formation, South Africa; Forest Sandstone Formation, Zimbabwe
Holotype- (QG 1) (2.15 m, 13 kg) (robust adult) (skull ~210 mm), fifth
dorsal vertebra (28 mm), sixth dorsal vertebra (28 mm), seventh dorsal vertebra
(29 mm), eighth dorsal vertebra (29 mm), ninth dorsal vertebra (29 mm), tenth
dorsal vertebra (32 mm), eleventh dorsal vertebra (31 mm), twelfth dorsal vertebra
(29 mm), thirteenth dorsal vertebra (29 mm), twelve dorsal ribs (sixth 141 mm),
gastralia, sacrum (123 mm), first caudal vertebra (24 mm), second-ninth caudal
vertebrae, tenth caudal vertebra (27 mm), eleventh to nineteenth caudal vertebra,
twenty-second caudal vertebra (26 mm), twenty-third to twenty-fifth caudal vertebra,
twenty-sixth caudal vertebra (27 mm), twenty-seventh caudal vertebra (29 mm),
twenty-eighth caudal vertebra (31 mm), twenty-ninth caudal vertebra, thirtieth
caudal vertebra (26 mm), thirty-first to thirty-eighth caudal vertebra, thirty-ninth
caudal vertebra (22 mm), fortieth caudal vertebra, thirty-six chevrons, scapulocoracoid
(130 mm), humerus (100 mm), radius (61 mm), ulna (77 mm), six carpals, metacarpal
I (12 mm), phalanx I-1 (20 mm), manual ungual I (18 mm), metacarpal II (26 mm),
phalanx II-1 (13 mm), phalanx II-2 (17 mm), manual ungual II (19 mm), metacarpal
III (26 mm), phalanx III-1 (10 mm), phalanx III-2 (9 mm), phalanx III-3 (12
mm), manual ungual III, metacarpal IV (17 mm), phalanx IV-1 (5 mm), ilium (148
mm), pubis (204 mm), ischium (130 mm), femur (208 mm), tibia (223 mm), fibula
(208 mm), astragalocalcaneum (29 mm wide), distal tarsal IV, metatarsal I (30
mm), phalanx I-1 (17 mm), pedal ungual I (14 mm), metatarsal II (119 mm), phalanx
II-1 (33 mm), phalanx II-2 (25 mm), pedal ungual II (22 mm), metatarsal III
(132 mm), phalanx III-1 (37 mm), phalanx III-2 (29 mm), phalanx III-3 (24 mm),
pedal ungual III (22 mm), metatarsal IV (117 mm), phalanx IV-1 (20 mm), phalanx
IV-2 (17 mm), phalanx IV-3 (15 mm), phalanx IV-4 (12 mm), pedal ungual IV (18
mm), metatarsal V (47 mm)
Referred- (BPI coll.) (at least eight individuals) cervical vertebra,
dorsal vertebra, distal femora, proximal tibia, pedal fragments (Raath 1980)
(QG 3A) (robust) femur (Raath, 1990)
(QG 76) (gracile) femur (201 mm) (Raath, 1990)
(QG 164) pes (Raath, 1977)
(QG 165) maxilla, nasals, lacrimal, jugal, quadratojugal, palatine, ectopterygoid,
pterygoid, hyoids, postcrania (Bristowe and Raath, 2004)
(QG 169) seventh cervical vertebra, eighth cervical vertebra, ninth cervical
vertebra, tenth cervical vertebra, first dorsal vertebra, second dorsal vertebra,
third dorsal vertebra, fourth dorsal vertebra (Raath, 1977)
(QG 170) eighth cervical vertebra (Raath, 1977)
(QG 171) second dorsal vertebra (Raath, 1977)
(QG 172) sixth cervical vertebra, seventh cervical vertebra, eighth cervical
vertebra (Raath, 1977)
(QG 173) cervical ribs (Raath, 1977)
(QG 175) third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra
(Raath, 1977)
(QG 176) atlas, axis, third cervical vertebra (Raath, 1977)
(QG 177) second cervical vertebra, third cervical vertebra, fourth cervical
vertebra (Raath, 1977)
(QG 179) third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra
(Raath, 1977)
(QG 180) second dorsal vertebra (Raath, 1977)
(QG 193) maxilla, nasal, lacrimal, prefrontal, jugal, frontals, parietals, postorbital,
squamosal, quadratojugal, braincase, mandible, scapulocoracoid, two furculae
(Raath, 1977; Tykoski et al., 2002)
(QG 194) maxilla, nasals, squamosal, quadratojugal, quadrate, braincase, endocranium,
ectopterygoid, pterygoid (Raath, 1977)
(QG 195) braincase, endocranium (Raath, 1977)
(QG 196) braincase, endocranium (Raath, 1977)
(QG 197) braincase, endocranium (Raath, 1977)
(QG 202) premaxilla, maxilla, nasal, dentaries, dentary teeth (Raath, 1977)
(QG 206) maxilla (Raath, 1977)
(QG 208) maxilla, sclerotic plates, mandible (Raath, 1977)
(QG 209) maxilla (Raath, 1977)
(QG 210) maxilla (Raath, 1977)
(QG 213) maxilla (Raath, 1977)
(QG 230) lacrimal (Raath, 1977)
(QG 234) lacrimal (Raath, 1977)
(QG 235) quadratojugal, quadrate, palatine, ectopterygoid, pterygoid (Raath,
1977)
(QG 241) palatine (Raath, 1977)
(QG 244) furcula (Raath, 1977; Tykoski et al., 2002)
(QG 245) premaxilla, premaxillary teeth (Raath, 1977)
(QG 246) premaxilla, premaxillary teeth (Raath, 1977)
(QG 249) premaxilla (Raath, 1977)
(QG 254) premaxilla (Raath, 1977)
(QG 263) pterygoid (Raath, 1977)
(QG 265) pterygoid (Raath, 1977)
(QG 278) jugal, lacrimal, prefrontal (Raath, 1977)
(QG 287) postorbital (Raath, 1977)
(QG 303) dentary (Raath, 1977)
(QG 305) dentary (Raath, 1977)
(QG 307) mandible (Raath, 1977)
(QG 406) fifth dorsal vertebra (Raath, 1977)
(QG 408) seventh dorsal vertebra (Raath, 1977)
(QG 514) (robust) humerus (Raath, 1990)
(QG 514b) (robust) ulna (Raath, 1990)
(QG 545) (gracile) humerus (Raath, 1990)
(QG 563) ulna
(QG 568) (gracile) ulna (Raath, 1990)
(QG 573) (robust) metacarpus (Raath, 1990)
(QG 577) (gracile) distal radius, distal ulna, manus (Raath, 1990)
(QG 687) distal carpal (Raath, 1977)
(QG 691) (gracile) pelvis, femur (142 mm), tibia, fibula, astragalus, calcaneum
(Raath, 1990)
(QG 713) (gracile) femur (Raath, 1990)
(QG 715) (gracile) femur (Raath, 1990)
(QG 716) femur (Raath, 1990)
(QG 717) (gracile) femur (Raath, 1990)
(QG 725) (robust) femur (Raath, 1990)
(QG 726) (robust) femur (Raath, 1990)
(QG 727) (robust) femur (Raath, 1990)
(QG 729) (robust) femur (Raath, 1990)
(QG 731) (robust) femur (189 mm) (Raath, 1990)
(QG 732) (robust) femur (Raath, 1990)
(QG 733) (robust) femur (Raath, 1990)
(QG 738) (gracile) femur (192 mm) (Raath, 1990)
(QG 739) (gracile) femur (Raath, 1990)
(QG 740) (gracile) femur (Raath, 1990)
(QG 742) (gracile) femur (Raath, 1990)
(QG 744) (gracile) femur (Raath, 1990)
(QG 745) (gracile) femur (Raath, 1990)
(QG 753) (robust) femur (Raath, 1990)
(QG 754) (robust) femur (186 mm) (Raath, 1990)
(QG 755) (robust) femur (185 mm) (Raath, 1990)
(QG 760) (robust) femur (Raath, 1990)
(QG 768) (gracile) tibia, fibula, astragalus, calcaneum, distal tarsal III,
distal tarsal IV, metatarsus (Raath, 1990)
(QG 786) (gracile) astragalus, calcaneum (Raath, 1990)
(QG 816) astragalus, calcaneum (Raath, 1977)
(QG 846) distal tarsal III (Raath, 1977)
Diagnosis- (modified from Tykoski and Rowe, 2004) differs from Coelophysis
bauri in the pit at the base of the nasal process of the premaxilla; Differs
from "Megapnosaurus" kayentakatae in promaxillary fenestra
absent; nasal crests absent; frontals not separated by midline anterior extension
of parietals; anterior astragalar surface flat.
(after Ezcurra, 2006) reduced distal medial condyle on metacarpal I.
Comments- Munyikwa and Raath (1999) described a partial skull as Syntarsus,
which was tentatively reidentified as Dracovenator by Yates (2005). Bristowe
and Raath (2004) used a partially articulated juvenile skull to show the nasal
fenestra identified by Raath (1977) was in fact closed in life, Raath articulated
the palatine backwards, and that his reconstruction of the lacrimal-jugal articulation
is inaccurate. They also confirmed the hyoids identified by Raath were furculae.
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