= Tetanurae sensu Sereno, 1998
Definition- (Passer domesticus <- Torvosaurus tanneri) (modified)
Reference- Sereno, 1998. A rationale for phylogenetic definitions, with application
to the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie
und Pal�ontologie Abhandlungen. 210(1), 41-83.
Avetheropoda Paul, 1988
Definition- (Allosaurus fragilis + Passer domesticus) (Holtz
et al., 2004; modified from Padian et al., 1999; modified from Currie and Padian,
1997)
= Dinoaves Bakker, 1986
= Tetanurae sensu Novas, 1992
Definition- (Allosaurus fragilis + Passer domesticus) (modified)
= Neotetanurae Sereno, Wilson, Larsson, Duthell and Sues, 1994
Definition- (Allosaurus fragilis + Passer domesticus) (Allain
et al., 2012; modified from Sereno, 1998)
Other definitions- (Sinraptor dongi + Carcharodontosaurus saharicus
+ Allosaurus fragilis + Passer domesticus) (Sereno, in press)
= Neotetanurae sensu Sereno, in press
Definition- (Sinraptor dongi + Carcharodontosaurus saharicus +
Allosaurus fragilis + Passer domesticus)
= Euavetheropoda Sorkin, 2015
Comments- Bakker (1986) used the name Dinoaves in a phylogenetic tree
for a clade containing allosaurs and what are now considered coelurosaurs, but
not ceratosaurs or podokesaurs (coelophysoids). It has not been used since and
could equally well apply to Tetanurae. Paul (1988) first used Avetheropoda as
an order containing allosaurids (including carcharodontosaurids) and all taxa
now placed in Coelurosauria, but not metriacanthosaurines (sinraptorids), megalosauroids
and more basal taxa. It was too largely ignored until Holtz (1994) used it for
an equivalent clade. Sereno proposed Neotetanurae for the same clade that year
and both names have since received equally frequent use (e.g. 54 Google Scholar
hits for Avetheropoda vs. 56 for Neotetanurae as of 1-15-2012). Sorkin (2015)
proposed the name Euavetheropoda in an abstract, for a clade containing Monolophosaurus,
acrocanthosaurids, allosaurids and coelurosaurs. It is uncertain if this is
based on an analysis, and such a clade excluding metriacanthosaurids, Neovenator,
carcharodontosaurines and megaraptorans has not been recovered in other studies.
Sereno's newest (in press) definition of Neotetanurae differs from the original
(Sereno, 1998) by adding Sinraptor and Carcharodontosaurus as
internal specifiers. I suppose it would preserve content better if sinraptorids
or carcharodontosaurids end up just basal to carnosaurs + coelurosaurs (Paul,
1988; Coria and Salgado, 1995; Longrich, 2001; Paul, 2002). However, if carcharodontosaurids
are ceratosaurs (Bonaparte et al., 1990) or sinraptorids are megalosaurids (Kurzanov,
1989), the original intent of Neotetanurae would be lost. The latter two situations
seem less likely than the former, so Sereno's redefinition may be more advantageous
than not.
While found in nearly every analysis, recent work suggests Avetheropoda as currently
conceived may not exist, with megalosauroids actually being carnosaurs so that
Avetheropoda includes the current content of Orionides. This can be found in
Rauhut (2003) and Cau's unpublished work, while Carrano et al. (2012) find it
in their trees with only two more steps.
References- Bakker, 1986. The Dinosaur Heresies. Kensington, New York.
481 pp.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Kurzanov, 1989. O proiskhozhdenii i evolyutsii infraotryada dinozavrov Carnosauria
[Concerning the origin and evolution of the dinosaur infraorder Carnosauria].
Paleontologicheskiy Zhurnal. 1989(4), 3-14.
Bonaparte, Novas and Coria, 1990. Carnotaurus sastrei Bonaparte, the
horned, lightly built carnosaur from the Middle Cretaceous of Patagonia. Natural
History Museum of Los Angeles County Contributions in Science. 416, 41 pp.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
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.
Sereno, Wilson, Larsson, Dutheil and Sues, 1994. Early Cretaceous dinosaurs
from the Sahara. Science. 266, 267-271.
Coria and Salgado, 1995. A new giant carnivorous dinosaur from the Cretaceous
of Patagonia. Nature. 377, 224-226.
Currie and Padian, 1997. Avetheropoda. In Currie and Padian (eds.). Encyclopedia
of Dinosaurs. Academic Press, New York. 39.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210(1), 41-83.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature
of the major taxonomic categories of the carnivorous Dinosauria (Theropoda).
Journal of Vertebrate Paleontology. 19(1), 69-80.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD Thesis. University of Bristol. 440 pp.
Longrich, 2001. Secondarily flightless maniraptoran theropods? Journal of Vertebrate
Paleontology. 21(3), 74A.
Paul, 2002. Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs
and Birds. Johns Hopkins University Press, Baltimore. 472 pp.
Holtz, Molnar and Currie, 2004. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 71-110.
Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian
spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften.
99(5), 369-377.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Sorkin, 2015. A re-evaluation of several character states in non-coelurosaurian
Tetanurae (Dinosauria: Theropoda) with implications for phylogeny of basal tetanurans.
Journal of Vertebrate Paleontology. Program and Abstracts 2015, 217.
Datanglong Mo, Zhou, Li,
Huang and Cao, 2014
D. guangxiensis Mo, Zhou, Li, Huang and Cao, 2014
Aptian?, Early Cretaceous
Xinlong Formation, Guangxi, China
Holotype- (GMG 00001) (adult) partial thirteenth dorsal vertebra (~130
mm), incomplete synsacrum (94, 78, 93, 113, 106 mm), incomplete first caudal
vertebra (~96 mm), incomplete second caudal vertebra, incomplete first chevron,
incomplete ilium (~700 mm), pubic fragment, proximal ischium
Diagnosis- (after Mo et al., 2014) thirteenth dorsal vertebra with teardrop-shaped
foramen bordered by centrum, anterior centroparapophyseal and posterior centrodiapophyseal
laminae; thirteenth dorsal vertebra with well developed horizontal prezygoparapophyseal
lamina; thirteenth dorsal vertebra with parapophysis projecting further laterally
than diapophysis; brevis fossa shallow with short, ridge-like medial blade;
iliac pubic peduncle with posteroventrally expanded margin.
Comments- The holotype was discovered in 2011. Mo et al. (2014) included
it in Carrano et al.'s tetanurine analysis and found it to be a carcharodontosaurid
outside Megaraptora and Acrocanthosaurus+Shaochilong+Carcharodontosaurinae.
Cau (online, 2014) noted this analysis only includes a few coelurosaurs, and
found that Datanglong emerges as a compsognathid-grade coelurosaur. If
more coelurosaurs are added to the Carrano tetanurine dataset, I find both positions
to be equally parsimonious. It is thus retained as Avetheropoda incertae sedis
here pending further study. Samathi and Chanthasit (2017) recovered Datanglong as a megaraptoran using Novas et al.'s tetanurine analysis, "sharing the pneumaticity of the ilium with other megaraptorans."
References- Cau, online 2014. http://theropoda.blogspot.com/2014/08/datanglong-un-nuovo-carcharodontosauria.html
Mo, Zhou, Li, Huang and Cao, 2014. A new Carcharodontosauria (Theropoda) from
the Early Cretaceous of Guangxi, Southern China. Acta Geologica Sinica (English
Edition). 88(4), 1051-1059.
Samathi and Chanthasit, 2017. Two new basal Megaraptora
(Dinosauria: Theropoda) from the Early Cretaceous of Thailand with
comments on the phylogenetic position of Siamotyrannus and Datanglong. Journal of Vertebrate Paleontology. Program and Abstracts, 188.
Lourinhanosaurus Mateus,
1998
L. antunesi Mateus, 1998
= Allosaurus antunesi (Mateus, 1998) Paul, 2010
Late Kimmeridgian-Early Tithonian, Late Jurassic
Sobral Formation, Portugal
Holotype- (ML 370) (4 m) cervical vertebrae, dorsal vertebrae, sacral
vertebrae, caudal vertebrae, chevrons, ilia, partial pubes, partial ischia,
partial femora, tibia, fibula, proximal metatarsal, 32 gastroliths
Referred- (ML 565) adult teeth, ~300 embryonic elements including maxilla, four teeth,
vertebrae, scapulae, ilium, femora, tibiae and metatarsi, ~100 eggs, nest (Mateus, 1997)
(ML 1194) eggshells (Ribeiro et al., 2013)
distal caudal vertebra (Mateus, pers. comm, 2002)
Diagnosis- (after Mateus, 1998) all vertebral centra are longer than
tall; proximal caudal neural spines with well-developed spike-like anterior
process; pubic blade perforated by large vertical ellipsoidal foramen; anterior
trochanter well separated from the main body axis of the femur in lateral view.
(after Carrano et al., 2012) medial condyle of tibia half the transverse width
of fibular condyle.
Comments- The specimen had 32 gastroliths and the enveloping sediment
preserved the negative imprint of 3 additional gastroliths. The maximum observed
gastrolith length is 22 millimetres. Near the pebbles there were three small
bone fragments that seem to be food remains. The gastroliths have been found
in the rib cage below the eleventh dorsal vertebra. The high number, concentration
and relative size of the gastroliths suggest that they belong to this specimen,
and that they had not been swallowed when eating other dinosaur's stomach. Mateus
(pers. comm., 2002) refers one distal caudal previously referred to Megalosaurus
insignis (Lapparent and Zbyszewski, 1957) to Lourinhanosaurus.
Antunes and Mateus (2003) referred "a femur (ML 555) at Porto das
Barcas" to the taxon, but Malafaia et al. (2020) later stated it "does
not share any diagnostic characters with the holotype; ML 555 has a
femoral head that projects dorsomedially, which is a feature that has
been interpreted as a synapomorphy of Carcharodontosauria." It is
here provisionally referred to contemporaneous carcahrodontosaurid Lusovenator.
Eggshells belong to Preprismatoolithus.
Phylogenetic relationships- Though Mateus (1998) originally referred this taxon to
Allosauroidea and Holtz et al. (2004) later found it to be a carnosaur, Allain
(2001) recovered it as a megalosaurid in his unpublished analysis while Mateus
et al. (2006) refer it to Eustreptospondylidae without explanation. Benson (2008,
2010) found it to be a sinraptorid, and in a more extensive analysis (Benson
et al., 2010) placed it sister to Streptospondylus
in that clade. Carrano et al. (2012) later found it to be a basal
coelurosaur, though only two more steps were needed to make it a basal
carnosaur or sister to Avetheropoda, so neither option is unlikely.
Hartman et al. (2019) recovered it as intermediate between
megalosauroids and sinraptorids, so a position close to the base of
Avetheropoda is advocated here.
References- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal.
M�moires du Service g�ologique du Portugal. 2, 1-63.
Mateus,
1997. Eggs, nest and embryos of theropod dinosaur in Upper
Jurassic level of Lourinh�, Portugal. XIII Encuentro de Jovenes
Investigadores Ponencias. Cuadernos de I.N.I.C.E.. 74-75, 215-217.
Mateus, Mateus, Antunes, Mateus, Taquet, Ribeiro and Manuppella, 1997. Couvee,
oeufs et embryons d'un Dinosaure Theropode du Jurassique de Lourinha (Portugal).
C.R Acad. Sci. Paris, Sciences de la terre et des plan�tes, 325: 71-78.
Mateus, 1998. Lourinhanosaurus antunesi, a new Upper Jurassic Allosauroid
(Dinosauria: Theropoda) from Lourinh� (Portugal). Mem�rias da
Academia de Ci�ncias de Lisboa. 37: 111-124.
Mateus, Mateus, Antunes, Mateus, Taquet, Ribeiro and Manuppella, 1998. Upper
Jurassic theropod dinosaur embryos from Lourinh� (Portugal). Mem�rias
da Academia de Ci�ncias de Lisboa. 37: 101-110.
Mateus, Taquet, Antunes, Mateus and Ribeiro, 1998. Theropod dinosaur nest from
Lourinha, Portugal. Journal of Vertebrate Paleontology, 18(3) 61A.
Allain, 2001. The phylogenetic relationships of Megalosauridae within basal
tetanurine theropods. Journal of Vertebrate Paleontology. 22(3), 31A.
Mateus, Antunes and Taquet, 2001. Dinosaur ontogeny: The case of Lourinhanosaurus
(Late Jurassic, Portugal). Journal of Vertebrate Paleontology, 21 (Suppl. 3):
78A.
Ricqles, Mateus, Antunes and Taquet, 2001. Histomorphogenesis of embryos of
Upper Jurassic Theropods from Lourinh� (Portugal). Comptes rendus de
l'Acad�mie des sciences - S�rie IIa - Sciences de la Terre et
des plan�tes. 332(10): 647-656.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle d�couverte
et r�vision syst�matique: Implications phylog�n�tiques
et pal�obiog�ographiques. Unpublished thesis. 329 pp.
Antunes and Mateus, 2003. Dinosaurs of Portugal. Comptes Rendus Palevol. 2(1), 77-95.
Cunha, Mateus and Antunes, 2004, The sedimentology of the Paimogo dinosaur nest
site (Portugal, Upper Jurassic): Abstract Book of the IAS [International Association
of Sedimentologists] 23rd Meeting, Coimbra, Portugal, p. 93.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska.
The Dinosauria Second Edition. University of California Press. 861 pp.
Benson, 2008. A new theropod phylogeny focussing on basal tetanurans, and its
implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism.
Journal of Vertebrate Paleontology. 51A.
Castanhinha, Araujo and Mateus, 2009. Dinosaur eggshell and embryo localities
in Lourinha Formation, Late Jurassic, Portugal. Journal of Vertebrate Paleontology.
29(3), 76A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. 158(4), 882-935.
Benson, Brusatte and Carrano, 2010. A new clade of large-bodied predatory dinosaurs
(Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97, 71-78.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Mateus, Carrano and Taquet, 2012. Osteology of the embryonic theropods from
the Late Jurassic of Paimogo, Portugal. Journal of Vertebrate Paleontology.
Program and Abstracts 2012, 137.
Ribeiro, Holwerda and Mateus, 2013. Theropod egg sites from the Lourinha Formation,
Portugal. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 198.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Mateus, Antunes, Taquet and Ricql�s, in preparation. The osteology of
the embryos of the theropod dinosaur Lourinhanosaurus antunesi from Portugal.
"Tyrannosaurus"
"lanpingensis" Ye, 1975
= Tyrannosaurus "lanpingi" Zhao, 1986
= Tarbosaurus "lanpinensis" (Ye, 1975) Azuma 1991
Valanginain-Barremian, Early Cretaceous
Jingxing (=Chingshong) Formation, Yunnan, China
Material- (IVPP coll.) partial tooth (FABL ~32 mm)
Comments- Ye (often cited as Yeh) first mentioned Tyrannosaurus lanpingensis
in 1975 as a new species from the IVPP that had not been published. Zhao (1986)
provides a photo of the basal half of a tooth, crediting the name Tyrannosaurus
"lanpingi" to himself (as Chao). In the plate's caption, he lists
it as being from the Early Cretaceous Jingxing Formation of Lanping, Yunnan.
Azuma (1991) listed "Tarbosaurus lanpinensis(?)" as a tyrannosaurid
from the Tugulu Group of China, citing Dong (1992, which was in press at the
time). Dong (1992) discusses it as Tyrannosaurus lanpingensis, mistakenly
attributing the name to Zhao, 1983. In Appendix A, Dong lists it as a tyrannosaurid
named Tarbosaurus (Tyrannosaurus) lanpinensis (no doubt
a misspelling, which Azuma's was based on), and mistakenly attributes it to
Chao, 1975. In Appendix C, it is Tarbosaurus (Tyrannosaurus) lanpingensis.
Dong states it is from the Chingshong Formation of Lanping County in Yunnan,
which he attributes to Tithonian-Neocomian age based on bivalves. He says the
species is based on a single tooth without diagnosis or drawing, so is a nomen
nudum, but Zhao (1986) did illustrate it. Ford and Chure misspelled it as Tyrannosaurus
langingi, credited to Zhao (1986), in an unpublished list based on their
2001 SVP abstract. They have it as from the Upper Chingshing Formation, which
they attribute to Berriasian-Hauterivian. Hurum and Sabath (2003) list "Tyrannosaurus
lanpingensis" as a tyrannosaurid, citing Ye. Azuma et al. (2006) lists
"Tyrannosaurus" lanpingensis from the Chingshong Formation.
The formation is almost always called the Jingxing Formation, and is agreed
by modern paleontologists to be Valanginian-Barremian (e.g. Sha, 2010).
The photo is the basal half of a tooth crown in labial view, and has a FABL
of ~32 mm. Only distal serrations are evident, which are small (8 per 5 mm)
and have apically angled blood grooves. The size and serration density fall
within the range of Tyrannosaurus rex, but also Carcharodontosaurus
saharicus. Angled blood grooves are known for tyrannosaurids and Carcharodontosaurus,
but also taxa like Fukuiraptor. Importantly, there seem to be prominent
enamel wrinkles along the distal carina. These are almost never found in tyrannosauroids,
but are known in derived carcharodontosaurids, Allosaurus and Fukuiraptor.
As "lanpingensis" is from the Early Cretaceous, it's more likely to
be a megaraptoran or carcharodontosaurid than an allosaurid or tyrannosaurine.
Both are clades known from large taxa (Chilantaisaurus; Carcharodontosaurus,
Giganotosaurus, Mapusaurus) and both are known from Asia (Chilantaisaurus,
Fukuiraptor; Shaochilong). Until more details are known, I recommend
placing "lanpingensis" in Avetheropoda indet..
References- Ye, 1975. Jurassic system. In Su (ed.). Mesozoic Redbeds
of Yunnan. Academia Sinica, Beijing. 11-30.
Zhao, 1986. The Cretaceous biota of China: Reptilia. in Hao, Su, Yu, Li, Li,
Wang, Qi, Guan, Hu, Liu, Yang, Ye, Shou, Zhang, et al.. The Cretaceous System
of China. Stratigraphy of China. 12, 67-73, plates XI, XII.
Azuma, 1991. Early Cretaceous dinosaur fauna from the Tetori Group Central Japan-
Research of dinosaurs from the Tetori Group (1). Professor Shizuka Miura Memorial
Volume. Fukui University, Fukui, Japan. 55-69.
Dong, 1992. Dinosaurian Faunas of China. Berlin, Heidelberg, New York, London,
Paris, Tokyo, Hong Kong: Springer-Verlag. 188 pp.
Ford and Chure, 2001. Ghost lineages and the paleogeographic and temporal distribution
of tyrannosaurids. Journal of Vertebrate Paleontology. 21(3), 50A-51A.
Hurum and Sabath, 2003. Giant theropod dinosaurs from Asia and North America:
Skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta
Palaeontologica Polonica. 48(2), 161-190.
Azuma, Li, Currie, Dong, Shibata and Lu, 2006. Dinosaur footprints from the
Lower Cretaceous of Inner Mongolia, China. Memior of the Fukui Prefectural Dinosaur
Museum. 5, 1-14.
Sha, 2010. Historical distribution patterns of trigonioidids (non-marine Cretaceous
bivalves) in Asia and their palaeogeographic significance. Proceedings of the
Royal Society B. 277(1679), 277-283.
"Unquillosauridae" Powell, 1986
Unquillosaurus Powell, 1979
U. ceibalii Powell, 1979
Campanian, Late Cretaceous
Los Blanquitos Formation, Salta, Argentina
Holotype- (PVL 3670-11) ilial fragment, pubis (514 mm)
Description- Novas and Agnolin (2004) determined the supposedly diagnostic
proximal sulcus noted by Powel (1979) doesn't exist, and is actually the pubic
peduncle of the ilium broken and displaced. This allowed them to identify a
ventrally concave pubic peduncle, and the angle between the anterior and ventral
edges supports opisthopuby. What's normally seen as the acetabular surface of
the pubis is the posterior part of a very long ilial peduncle, leaving a very
tiny space for the acetabulum, which is said to resemble maniraptoriformes.
These characters support placement in Maniraptora.
Comments- While the family Unquillosauridae was apparently named by Powell
in his thesis, the ICZN does not accept theses as valid for nomenclatural purposes,
so it is a nomen nudum.
In 1997, Ford suggested on the DML that Unquillosaurus was similar to
Unenlagia, which has gone so far as to prompt discussion of synonymy.
However, Unquillosaurus is not synonymous with Unenlagia. First,
I should note the pubis was described incorrectly, the medial side being lateral
and vice versa (as later determined by Carrano et al., 2012). The supposed "lateral
crest" is really the pubic apron, the "lateral" facets on the
pubic boot are really for the pubic symphysis. Compared to Unenlagia-
the pubis is more propubic; the ischial peduncle is longer; there was an obturator
notch; the shaft is anteroposteriorly thicker distally; the pubic boot projects
slightly anteriorly; there is a proximomedial sulcus; the ilial peduncle is
less transversly expanded; the pelvic canal is narrower; the proximal shaft
expands laterally; the distal end is expanded transversely, not compressed;
there is a gap in the symphysis proximal to the pubic boot. Also keep in mind
Unquillosaurus is from the Los Blanquitos Formation, while Unenlagia
is from the Rio Neuquen Formation.
Novas and Agnolin (2004) keep Powell's orientation, remarking on the "prominent
external longitudinal ridge" (=pubic apron?), odd medially convex pelvic
canal margins (concave if reversed) and lack of an apron or medial symphysis.
In addition, the medial end is described as "flattened and lacks marks
for the articulation with the opposite bone". I see no reason to doubt
the hypothesis they have the pubis reversed.
Novas and Agnolin refer to Maniraptora, while Carrano et al. suggest it is a
carcharodontosaurid. It is at least an avetheropod due to lacking an obturator
foramen.
References- Powell, 1979. Sobre una asociacion de dinosaurios y otras
evidencias de vertebrados del Cretacico superior de la region de La Candelaria,
Prov. de Salta, Argentina. Ameghiniana. 16, 191-204.
Powell, 1986. Revision de los titanosauridos de America del Sur Argentina. PhD
thesis. Universidad Nacional de Tucuman, Tucuman, Argentina. 340 pp.
Novas and Agnolin, 2004. Unquillosaurus ceibalii Powell, a giant maniraptoran
(Dinosauria, Theropoda) from the Late Cretaceous of Argentina. Revista del Museo
Argentino de Ciencias Naturales, nuevo serie. 6(1), 61-66.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed avetheropod (Turner, Hwang and Norell, 2007)
Berriasian-Barremian, Early Cretaceous
Huhteeg Svita, Mongolia
Holotype- (IGM coll.) postorbital (60 mm)
Comments- The heavily rugose texture and straight anterior process suggest
assignment to Carnosauria or Tyrannosauroidea.
Reference- Turner, Hwang and Norell, 2007. A small derived theropod from
Oosh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates. 3557,
27 pp.
unnamed possible avetheropod (Hu, 1963)
Late Valanginian-Early Albian, Early Cretaceous
Jehol Group, Liaoning, China
Material- (IVPP V2756) partial mid caudal vertebra, proximal tibia, distal
phalanx
Comments- Hu (1963) assigned this to Megalosauridae indet., but Carrano
et al. (2012) believed the small lateral condyle was more similar to allosaurians
and coelurosaurs. They stated it may belong to the latter clade. Additionally,
I note the distally restricted fibular crest is like Cristatusaurus, Torvosaurus,
allosaurians and coelurosaurs.
References- Hu, 1963. [The carnivorous dinosaurian remains from Fusin,
Liaoning]. Vertebrata PalAsiatica. 7, 174-176.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed possible avetheropod (Janensch, 1925)
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (MBR 3628; = St 233) ilium (537 mm)
Comments- Rauhut (2011) found this to be most similar to megalosauroids,
except for the cuppedicus fossa which is like some ceratosaurs and avetheropods.
It is not coelurosaurian however. Carrano et al. (2012) stated the transverse
pubic peduncle compression (~2 times long as wide) is like allosaurians and
suggested it was carcharodontosaurian based on the more vertical pubic peduncle
orientation, but the peduncle is only 1.78 times longer than wide and a ventrally
projecting pubic peduncle is shared with almost all averostrans in their matrix.
When entered into Carrano et al.'s matrix, MBR 3628 emerges as a non-allosauroid
avetheropod excluded from Proceratosaurus+Ornitholestes+Compsognathus.
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. 1(supp. 7), 1-99.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed avetheropod (Forster, Farke, McCartney, De Klerk and Ross, 2009)
Berriasian-Valanginian, Early Cretaceous
Kirkwood Formation, South Africa
Material- (AM 6041) proximal femur
Comments- Forster et al. (2009) referred this to non-avetheropod Tetanurae
because of its anteriorly angled head, but Carrano et al. (2012) noted this
is also present in some carnosaurs, and that the lack of an articular groove
and presence of an accessory trochanter were avetheropod characters. They thus
referred it to Carnosauria, but did not realize basal coelurosaurs like Tugulusaurus
have anteriorly angled heads as well.
References- De Klerk, Forster, Sampson, Chinsamy-Turan and Ross, 2000.
A new coelurosaurian dinosaur from the early Cretaceous of South Africa. Journal
of Vertebrate Paleontology. 20(2), 324-332.
Forster, Farke, McCartney, De Klerk and Ross, 2009. A "basal" tetanuran
from the Lower Cretaceous Kirkwood Formation of South Africa. Journal of Vertebrate
Paleontology. 29(1), 283-285.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed Avetheropoda (Benson, Rich, Vickers-Rich and Hall, 2012)
Early Aptian, Early Cretaceous
Wonthoggi Formation of the Strzelecki Group, Victoria, Australia
Material- (NMV P186057) incomplete ilium
(NMV P186327) partial coracoid
Reference- Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna
from southern Australia indicates high polar diversity and climate-driven dinosaur
provinciality. PLoS ONE. 7(5), e37122.
unnamed Avetheropoda (Brougham, Smith and Bell, 2019)
Early Cenomanian, Late Cretaceous
Griman Creek Formation, New South Wales, Australia
Material- (AM F105662) proximal femur
(AM F106525) central fragment
(LRF 3310) partial proximal caudal vertebra
....(LRF 3311) partial proximal caudal centrum
....(LRF 3312) ilial fragment
Comments- Brougham et al.
(2019) described several fragments as Avetheropoda indet.. They
state "LRF 3310-3312 presents a combination of characters that indicate
a probable phylogenetic position within Avetheropoda: camellae in the
caudal centra, a ventral keel on the anterior caudal centra and a pubic
peduncle approximately twice as long anteroposteriorly as
mediolaterally wide" and that "LRF 3310 and AM F106525 both have in
common a central convexity on the articular surface of the centrum, an
uncommon feature among theropods. The shared presence of this unusual
characteristic in both vertebrae, together with their relative
proximity to each other suggests that they may pertain to the same
taxon, or similar taxa." AM F105662 is reported to lack a
proximal articular groove as in avetheropods but lacks a concave
proximal edge unlike many coelurosaurs.
Reference- Brougham, Smith and
Bell, 2019. New theropod (Tetanurae: Avetheropoda) material from the
'mid'-Cretaceous Griman Greek Formation at Lightning Ridge, New South
Wales, Australia. Royal Society Open Science. 6, 180826.
Carnosauria Huene, 1920
Definition- (Allosaurus fragilis <- Passer domesticus)
(modified from Holtz et al., 2004; modified from Holtz and Padian, 1995)
Other definition- (Megalosaurus bucklandii, Allosaurus fragilis <- Passer domesticus) (modified from Rauhut and Pol, 2019)
= Allosauroidea sensu Sereno, 1998
Definition- (Allosaurus fragilis <- Passer domesticus) (modified)
= "Yangchuanosauria" Longrich, 2002
Definition- (Yangchuanosaurus shangyouensis <- Passer domesticus)
(modified from Longrich, 2002)
= Allosauroidea sensu Rauhut and Pol, 2019
Definition- (Allosaurus fragilis <- Megalosaurus bucklandii, Passer domesticus)
References- Huene, 1920. Bemerkungen zur Systematik und Stammesgeschichte
einiger Reptilien [Remarks on the systematics and phylogeny of some reptiles].
Zeitschrift f�r Induktive Abstammungs und Vererbungslehre. 22, 209-212.
Holtz and Padian, 1995. Definition and diagnosis of Theropoda and related taxa.
Journal of Vertebrate Paleontology. 15(3), 35A.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210(1), 41-83.
Longrich, 2002. Systematics of Sinosauropteryx. Journal of Vertebrate
Paleontology. 22(3), 80A.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle
Jurassic Ca�ad�n Asfalto Formation of Argentina highlights phylogenetic
uncertainty in tetanuran theropod dinosaurs. Scientific Reports.
9:18826.
"Asfaltovenator" Rauhut and Pol, 2019
"A. vialidadi" Rauhut and Pol, 2019
Late Toarcian-Bajocian, Early-Middle Jurassic
Ca�ad�n Asfalto Formation, Chubut, Argentina
Material- (MPEF PV 3440)
(~7-8 m) skull (~750-800 mm), mandible, ten cervical vertebrae,
cervical ribs, thirteen dorsal vertebrae, dorsal ribs, first sacral
vertebra, scapula, coracoid, humeri (335, 343 mm), radii (~200, ~205
mm), ulnae (250, 232 mm), radiale, intermedium, distal carpal I, distal
carpal II, metacarpal I (59 mm), phalanx I-1 (112 mm), manual ungual I
(~97 mm), metacarpal II (109 mm), phalanx II-1 (77 mm), phalanx II-2
(83 mm), manual unguals II (77 mm), metacarpal III (95 mm), phalanx
III-1 (32 mm), phalanx III-2 (~28 mm), phalanx III-3 (39 mm), manual
ungual III (~53 mm), distal pubes, distal femur, proximal tibia,
proximal fibula, distal tarsal IV, metatarsal II, metatarsal III,
phalanx III-1, phalanx III-2, metatarsal IV, phalanx IV-1, phalanx
IV-3, phalanx IV-4
Diagnosis- (after Rauhut and
Pol, 2019) premaxillary teeth with well-developed distal, but only
minute mesial serrations; postorbital with small cornual dorsal
process; exoccipital with pronounced horizontal ridges between
paroccipital processes and foramen magnum; antarticular in mandible;
platycoelous cervical vertebrae; neural spines of third and fourth
cervical vertebrae triangular and backswept; anterior cervical
epipophyses tab-like and elongated; mid cervical vertebrae with median
pit between parapophyses ventrally; ventral keel absent in posterior
cervical and poorly developed in anterior dorsal vertebrae;
well-developed paradiapohyseal lamina in middle and posterior dorsal
vertebrae; eleventh and twelfth dorsals with small additional anterior
centrodiapophyseal lamina; metacarpus broader than long; manual
digit III significantly more slender and shorter than digits I and II.
Comments- Pol and Rauhut (2012) originally noted this as a
large partially articulated basal tetanurine. Rauhut and Diego (2012)
in an abstract reported the mix of characters supporting and arguing
against assignment to Tetanurae, Megalosauroidea and Carnosauria, with
a suggestion this could indicate carnosaurian megalosauroids or high
amounts of homoplasy. Rauhut and Pol (2019) named and described the
taxon, but this paper has no mention of ZooBank and as of January 23
2020
"Asfaltovenator" lacks an entry on the ZooBank webite. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Asfaltovenator vialidadi"
Rauhut and Pol, 2019 is a nomen nudum that may never be technically
valid
as its journal is not published physically.
Rauhut and Pol added "Asfaltovenator" to Carrano et al.'s tetanurine
analysis and recovered it sister to allosauroids in Carnosauria, with
piatnitzkysaurids, Xuanhanosaurus, megalosaurids, spinosaurids and Monolophosaurus successively more basal non-allosauroid carnosaurs. Forcing "Asfaltovenator" in Megalosauroidea takes 4 steps
References- Pol and Rauhut, 2012. A Middle Jurassic abelisaurid from Patagonia
and the early diversification of theropod dinosaurs. Proceedings of the Royal
Society B. 279(1741), 3170-3175.
Rauhut and Diego, 2012. A new basal tetanuran theropod from the Early Middle
Jurassic of Patagonia, Argentina. Journal of Vertebrate Paleontology. Program
and Abstracts 2012, 160.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle
Jurassic Ca�ad�n Asfalto Formation of Argentina highlights phylogenetic
uncertainty in tetanuran theropod dinosaurs. Scientific Reports.
9:18826.
Allosauroidea Marsh, 1878 vide Currie and Zhao,
1994
Definition- (Allosaurus fragilis + Sinraptor dongi) (Holtz
et al., 2004; modified from Padian and Hutchinson, 1997)
Other definitions- (Allosaurus fragilis <- Passer domesticus)
(Brusatte and Sereno, 2008; modified from Sereno, 1998)
(Allosaurus fragilis <- Megalosaurus bucklandii, Passer domesticus) (Rauhut and Pol, 2019)
Comments- Note that while volume 30(10) of the Canadian Journal of Earth
Sciences lists its date as October 1993, it was not published until February
or March of 1994.
References- Marsh, 1878. Notice of new dinosaurian reptiles. American
Journal of Science and Arts. 15, 241-244.
Currie and Zhao, 1994. A new carnosaur (Dinosauria, Theropoda) from the Jurassic
of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences.
30(10), 2037-2081.
Padian and Hutchinson, 1997. Allosauroidea. In Currie and Padian (eds.). Encyclopedia
of Dinosaurs. Academic Press, New York. 6-9.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210(1), 41-83.
Holtz, Molnar and Currie, 2004. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 71-110.
Brusatte and Sereno, 2007. Phylogeny of Allosauroidea (Dinosauria: Theropoda:
New analysis, comparisons, and sources of disagreement. Journal of Vertebrate
Paleontology. 27(3), 54A.
Brusatte and Sereno, 2008. Phylogeny of Allosauroidea (Dinosauria: Theropoda):
Comparative analysis and resolution. Journal of Systematic Palaeontology. 6(2),
155-182.
Bates, Benson and Peter, 2010. The evolution of body size, stance and gait in
Allosauroidea (Dinosauria: Theropoda). Journal of Vertebrate Paleontology. Program
and Abstracts 2010, 58A.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle
Jurassic Ca�ad�n Asfalto Formation of Argentina highlights phylogenetic
uncertainty in tetanuran theropod dinosaurs. Scientific Reports.
9:18826.
Erectopodidae Huene, 1932
Erectopus Huene, 1923
E. superbus (Sauvage, 1882a) Huene, 1923
= Megalosaurus superbus Sauvage, 1882a
= gen. indet. superbus (Sauvage, 1882a) Huene, 1932
= Erectopus sauvagei Huene, 1932
Early Albian, Early Cretaceous
Sables verts, Meuse, France
Lectotype- (MNHN 2001-4) anterior maxilla
'Plastotype'- (MNHN 2001-4; holotype of Erectopus sauvagei) incomplete
phalanx I-1 (55 mm), partial manual ungual I, incomplete phalanx II-I (28 mm),
phalanx II-2 (28 mm), partial manual ungual II, distal metacarpal III, phalanx
III-1 (25 mm), proximal phalanx III-2, femur (480 mm), proximal tibia, distal
tibia, calcaneum, metatarsal II (230 mm)
Paralectotypes- (lost) teeth, dorsal centra (53, 56, 62, 65, 65 mm),
dorsal rib fragments, sacral fragment, proximal caudal vertebra (58 mm), distal
caudal vertebra (75 mm), neural spine fragments, distal radius(?), distal ulna(?),
metacarpal I (45 mm), phalanx I-1 (45 mm), metacarpal IV, partial ilium, proximal
fibulae, phalanx IV-? (30 mm)
Referred- (SV2) distal tibia (Buffetaut and Nori, 2012)
(SV3) incomplete metatarsal III (Buffetaut and Nori, 2012)
?(SV4) tooth (37.7x18.6x10.3 mm) (Buffetaut and Nori, 2012)
?(SV5) tooth (62.8x21.8x12.4 mm) (Buffetaut and Nori, 2012)
?(SV6) tooth (25x?x8.5 mm) (Buffetaut and Nori, 2012)
Comments- Barrois (1875) and Sauvage (1876) described a tooth from Louppy-le-Chateau
and other remains (two teeth and a centrum from Grandpre) as Megalosaurus.
Sauvage (1882a) later briefly described a partial skeleton which is the type
of his new species Megalosaurus superbus, associated with a larger distal
femur. He described the skeleton, distal femur and other remains later that
year (1882b), believing the previously known teeth from Louppy and Grandpre
to come from the same species. Huene (1923; and later in 1926a, b) separated
it from Megalosaurus as the new genus Erectopus superbus. Sauvage
did not designate any material as a holotype, and it is not certain it all derives
from one individual. In particular, the larger distal femur was found in the
same area, and the teeth were discovered before the partial skeleton in a potentially
different area. Huene (1932) incorrectly believed Sauvage based the name superbus
on the original teeth, and as he thought these were too large to go with the
postcrania, he separated the latter as Erectopus sauvagei. However, there
is no evidence Sauvage intended to base superbus on the teeth, and the
fact he waited until describing the postcrania in 1882 to name the species suggests
the opposite. Furthermore, Allain (2005) correctly notes the original teeth
match the preserved maxillary teeth in size. Huene retained the teeth as gen.
indet. superbus, which he believed was an allosaurid in contrast to Erectopus,
which he placed in a monotypic new family. Allain made the maxillary fragment
the lectotype of Erectopus superbus, as the rest of the material is lost
(though some elements are preserved as casts to form a plastotype). As the tooth
matches that on the lectotype, and the rest of the specimen is no more certainly
associated than the maxilla and tooth are with it, there is no reason to separate
the tooth from the postcrania taxonomically. Notably, the ICZN does not allow
a species to lack a genus, and the genus Erectopus must be attached to
the species superbus based on Huene (1923), regardless which elements
each name is based on. Erectopus sauvagei is thus an objective junior
synonym of Erectopus superbus.
Several elements have been reidentified since Sauvage's (1882a, b) description.
The posterior mandible is an anterior maxilla (Allain, 2005).The clavicle was
reidentified as a scapula (Huene, 1926a, b), but is a partial ilium (Chure,
2000). The "inferior" (=proximal?) radius is a distal tibia (Huene,
1926a, b). Huene (1926a, b) identified two supposed proximal metacarpals as
a distal radius and ulna, though as these were never illustrated this is uncertain.
Manual digits II-IV are I-III, and phalanges II-2 and II-3 are actually just
I-2 (the ungual). Similarly, phalanges III-3 and III-4 are just II-3 (the ungual).
The supposed manual phalanx III-1 of Sauvage and Chure (?-1 of Huene) is more
probably the other phalanx I-1 as it is too long for III-1, and too broad for
digit III. Sauvage identified an element as the lateralmost metacarpal, which
would be V in his reconstruction, but which is probably IV (considered a lateral
manual phalanx by Chure, 2000). The questionably referred manual phalanx of
Sauvage and pedal phalanx I-1 of Huene is metacarpal I (Molnar, 1990). The distal
fibulae are proximal fibulae (Huene, 1926a, b).
Not Erectopus- Several other European remains have been referred
to Erectopus or Megalosaurus superbus in the past. Sauvage (1882b)
thought the tooth from Louppy and two teeth from Grandpre described by Barrois
(1875) and Sauvage (1876) belonged to his new species Megalosaurus superbus.
A distal metatarsal and a proximal metapodial from Grandpre were also described,
the latter of which Huene (1926a, b) believed was a distal fibula. Sauvage also
referred a much larger distal femur from Louppy to Megalosaurus superbus,
though Huene (1926a, b) believed it to derive from a different taxon. Also figured
by Sauvage is a large pedal phalanx from Bar-le-Duc, while he mentions another
large pedal phalanx from an unstated locality.
Simionescu (1913) compared a tooth (UAIC (SCM1) 615) from the Valanginian Cernavoda
Formation of Romania to Megalosaurus superbus. Huene (1926) doubted it
was the same species due to age differences, but did continue to call it Erectopus
aff. superbus. It was recently redescribed by Csiki-Sava et al. (2016)
as a carcharodontosaurine.
Huene (1926a) referred a centrum from Grandpre described as Megalosaurus
by Sauvage (1876) to Erectopus superbus. Similarly, he referred a distal
femur from Blacourt described as dinosaurian by Sauvage (1876) to E. superbus,
but Chure (2000) determined it is crocodilian.
Stromer (1934) referred a proximal tibia, distal femur and distal tibia (IPHG
1912 VIII 78, 85 and 190) from the Cenomanian Baharija Formation of Egypt to
aff. Erectopus superbus, but none are similar to that genus.
Lapparent and Zbyszewski (1957) referred two tooth fragments from the Aptian
of Portugal to Megalosaurus superbus, but these are undiagnostic.
Relationships- Allain (2002a, b) found Erectopus to emerge as
a carnosaur in an unpublished phylogenetic analysis based on his thesis, which
was also the conclusion of his 2005 redescription of the material. This was
based solely on the interpretation of the distal tibia as indicating a well
developed posterior astragalar ascending process. However, a similar morphology
is also seen in the tibia of Poekilopleuron, which lacks a posterior
ascending process. Carrano et al. (2012) suggest Erectopus is a non-carcharodontosaurid
allosauroid, possibly a metriacanthosaurid, though they did not include it in
their analysis. When it is added to the matrix, it emerges as a non-allosaurian
carnosaur, and indeed can be a metriacanthosaurid. Note if it is a metriacanthosaurid,
Erectopodidae would have priority as the family name.
References- Barrois, 1875. Les reptiles du terrain Cr�tac�
du nord-est du Bassin de Paris. Bulletin scientifique, historique et litt�raire
du Nord. 6, 1-11.
Sauvage, 1876. Notes sur les reptiles fossiles no. 9. De la presence du type
dinosaurien dans le Gault du nord de la France. Bulletin de la Soci�t�
G�ologique de France. 4, 439-442.
Sauvage, 1882a. Sur les Reptiles trouv�s dans le gault de l'est de la
France [On the reptiles found in the Gault of eastern France]. Comptes Rendus
Hebdomadaires des Seances de l'Acad�mie des Sciences. 94, 1265-1266.
Sauvage, 1882b. Recherches sur les reptiles trouv�s dans le Gault de
l'est du bassin de Paris [Research on the reptiles found in the Gault of the
eastern Paris Basin]. M�moires de la Soci�t� G�ologique
de France, s�rie 3. 2(4), 1-42.
Simionescu, 1913. Megalosaurus aus der Unterkreide der Dobrogea. Centralblatt
f�r Mineralogie, Geologie und Pal�ontologie. 1913(20), 686-687.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of
the Geological Society of America. 34, 449-458.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
Huene. 1926b. On several known and unknown reptiles of the order Saurischia
from England and France. Annals and Magazine of Natural History.17, 473-489.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers in den W�sten
�gyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes Cenoman). 13.
Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche
Abteilung, Neue Folge. 22, 1-79.
Lapparent and Zbyszewski, 1957. [The dinosaurs of Portugal]. Services Geologiques
du Portugal. Memoire 2, 1-63.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California Press,
Berkeley, Los Angeles, Oxford. 306-317.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle d�couverte
et r�vision syst�matique: Implications phylog�n�tiques
et pal�obiog�ographiques. PhD thesis. Museum National d'Histoire
Naturelle Laboratoire de Paleontologie. 329 pp.
Allain, 2002b. The phylogenetic relationships of Megalosauridae within basal
tetanurine theropods (Dinosauria). Journal of Vertebrate Paleontology. 22(3),
31A.
Allain, 2005. The enigmatic theropod dinosaur Erectopus superbus (Sauvage,
1882) from the Lower Albian of Louppy-le-Ch'teau (Meuse, France). In Carpenter
(ed.). The Carnivorous Dinosaurs. Indiana University Press. 72-86.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Buffetaut and Nori, 2012. Dinosaur remains from the "Sables verts"
(Early Cretaceous, Albian) of the Eastern Paris Basin. In Godefroit (ed.). Bernissart
Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press.
362-377.
Csiki-Sava, Brusatte and Vasile, 2016. "Megalosaurus cf. superbus"
from southeastern Romania: The oldest known Cretaceous carcharodontosaurid (Dinosauria:
Theropoda) and its implications for earliest Cretaceous Europe-Gondwana connections.
Cretaceous Research. 60, 221-238.
Kelmayisaurus Dong, 1973
K. petrolicus Dong, 1973
Valanginian-Albian, Early Cretaceous
Lianmuqin Formation of the Tugulu Group, Xinjiang, China
Holotype- (IVPP V4022) maxillary fragment, quadrate fragment (lost),
incomplete dentary (523 mm)
Referred- ? distal pubes (Dong, 1992)
Early Cretaceous
Ejinhoro Formation, Inner Mongolia, China
Referred- ? (CCDP coll.) mandible (Dong, 1992)
Diagnosis- (after Brusatte et al., 2010) deeply inset and dorsally concave
accessory groove located anteriorly on the lateral surface of the dentary.
Comments- Generally viewed as an indeterminate theropod, Brusatte et
al. (2010) report Kelmayisaurus is diagnostic and shares characters with
megalosauroids, carcharodontosaurids and megaraptorans. They redescribed it
the following year as a basal carcharodontosaurid more derived than Neovenator
but less than Acrocanthosaurus and more derived taxa, but only one more
step was necessary to place it as a more basal carcharodontosaurid, megalosaurine
or basal coelurosaur. Adding it to the matrix of Caranno et al. (2012) results
in a less resolved placement however, as an allosaurian excluded from Megaraptora
and carcharodontosaurids as derived as Eocarcharia. Moving it to Megalosauroidea
or Coelurosauria still requires only a single additional step, so either relationship
is about equally probable.
Dong (1992) mentioned two referred specimens. One is from the Ejinhoro Formation
and may be possible to refer to Kelmayisaurus as the holotype also includes
a dentary. The other is a pair of distal pubes from the same locality as the
holotype, but cannot be compared with the latter.
References- Dong, 1973. Dinosaurs from Wuerho. Memoirs of the Institute
of Vertebrate Paleontology and Paleoanthropology Academica Sinica. 11, 45-52.
Dong, 1992. Dinosaurian Faunas of China. Ocean Press/Springer-Verlag, Beijing/Berlin.
188 pp.
Brusatte, Benson and Xu, 2010. The evolution of large-bodied theropod dinosaurs
during the Mesozoic in Asia. Journal of Iberian Geology. 36(2), 275-296.
Brusatte, Benson and Xu, 2012. A reassessment of Kelmayisaurus petrolicus,
a large theropod dinosaur from the Early Cretaceous of China. Acta Palaeontologica
Polonica. 57(1), 65-72.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Xuanhanosaurus Dong,
1984
X. qilixiaensis Dong, 1984
Bajocian, Middle Jurassic
Xuanhan 7 Mile Gorge, Xiashaximiao Formation, Sichuan, China
Holotype- (IVPP V6729) four anterior dorsal vertebrae, posterior dorsal
vertebra, dorsal neural arch, partial scapula, coracoid, humerus (265 mm), radius
(202 mm; proximal end lost), ulna (240 mm; lost), distal carpal I, distal carpal
II, distal carpal III, metacarpal I (52 mm), phalanx I-1 (84 mm), manual ungual
I (64 mm), metacarpal II (109 mm), phalanx II-1 (64 mm), metacarpal III (94
mm), phalanx III-1 (32 mm), phalanx III-2 (28 mm), metacarpal IV (50 mm)
Diagnosis- (after Rauhut, 2000) glenoid articular facet of humerus forms
a raised horizontal ridge that overhangs the humeral shaft posteriorly.
(after Carrano et al., 2012) dorsal neural spines transversely thick with gently
concave lateral surfaces.
Comments- The supposed sternum is part of the coracoid (Rauhut, 2000).
Benson (2008, 2010) and Holtz et al. (2004) recovered Xuanhanosaurus
as a non-orionidan tetanurine, and Rauhut (2000) also found it to be a non-coelurosaur
tetanurine. Benson found it to be sister to the Marshosaurus + Piatnitzkysaurus
+ Condorraptor clade, but this was poorly supported. More recently, Carrano
et al. (2012) recovered the genus as a metriacanthosaurid outside Yangchuanosaurus
and the derived metriacanthosaurine clade (Sinraptor, Siamotyrannus
and Metriacanthosaurus). Yet when their analysis is properly ordered,
it has a less exact position as a carnosaur outside Yangchuanosaurus,
the derived metriacanthosaurine clade and Allosauria. In Carrano et al.'s matrix,
it moves to Piatnitzkysauridae with only one more step, and outside Orionides
with only three more steps, so no suggested position is well supported.
References- Dong, 1984. A new theropod dinosaur from the Middle Jurassic
of Sichuan Basin. Vertebrata PalAsiatica. 22(3), 213-218.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Benson, 2008. A new theropod phylogeny focusing on basal tetanurans and its
implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism.
Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Xu, Han and Zhao, 2014. Homologies and homeotic transformation of the theropod
'semilunate' carpal. Nature Scientific Reports. 4, 6042.
unnamed probable allosauroid (Russell, 1996)
Albian, Early Cretaceous
Gres Rouges Infracenomaniens, Morocco
Material- (CMN 41865; bone taxon F) distal humerus (~385 mm, 156 mm wide)
Comments- Very similar to Xuanhanosaurus.
Reference- Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous
of the Tafilalt, Morocco. Bulletin du Museum national d'Histoire naturelle.
18, 349-402.
undescribed allosauroid (Kirkland, 2005)
Barremian, Early Cretaceous
Yellow Cat Member of Cedar Mountain Formation, Utah, US
Comments- Kirkland (2005) listed a "large carnosaurid perhaps related
to Utah’s state fossil, the Late Jurassic Allosaurus" as coming
from the Yellow Cat Member.
Reference- Kirkland, 2005. Utah’s newly recognized dinosaur record.
Utah Geological Survey: Survey Notes. 37(1), 1-5.
undescribed allosauroid (Judd, Irmis and Kirkland, 2013)
Early Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- cervical vertebra, partial sacrum, partial pelvis, femur
Comments- Judd et al. (2013) found this to be an allosauroid based on-
anterior pleurocoel of cervical vertebra anteroposteriorly elongate; cervical
parapophysis located in middle of centrum; no articular groove on proximal surface
of femoral head. However, it is similar to megalosaurids in having an oblique
ligament groove that does not extend past the posterior surface of the femoral
head.
Reference- Judd, Irmis and Kirkland, 2013. A new large-bodied theropod
dinosaur from the Lower Cretaceous Cedar Mountain Formation (Ruby Ranch Member)
in Central Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2013,
150.
unnamed possible allosauroid (Williston, 1902)
Early-Middle Albian, Early Cretaceous
Kansas, US
Material- centrum
References- Williston, 1902. Notes on some new or little-known extinct
Reptiles. Kansas University Science Bulletin. 1, 247-254.
Lane, 1946. A survey of the fossil vertebrates of Kansas Part III: The Reptiles.
Transactions of the Kansas Academy of Science. 49(3), 289-332.
unnamed Allosauroidea (Naish, 2003)
Valanginian, Early Cretaceous
Hastings Group, England
Material- (HASMG G.378; = HASTM GG98 of Benton and Spencer, 1995) proximal
tibia (~550 mm)
teeth (Austen et al., 2010)
Comments- The tibia differs from Neovenator, though it resembles
the latter and Allosaurus more than Fukuraptor or Sinraptor.
It may be referrable to Altispinax.
References- Benton and Spencer, 1995. Fossil Reptiles of Great Britain.
Chapman & Hall, London.
Naish, 2003. A definitive allosauroid (Dinosauria; Theropoda) from the Lower
Cretaceous of East Sussex. Proceedings of the Geologists' Association. 114,
319-326.
Austen, Brockhurst and Honeysett, 2010. Vertebrate fauna from Ashdown Brickworks,
Bexhill, East Sussex. Wealden News. 8, 13-23.
unnamed allosauroid (Benson, Brusatte, Hutt and Naish, 2009)
Barremian, Early Cretaceous
Wessex Formation, England
Material- (MIWG 6350) (~5.3 m) incomplete pubes, distal femur
Comments- Benson et al. (2009) identified this as a nonm-coelurosaurian
tetanurine, while Carrano et al. (2012) further indicated it was likely to be
a carnosaur due to the large pubic boot.
References- Benson, Brusatte, Hutt and Naish, 2009. A new large basal
tetanuran (Dinosauria: Theropoda) from the Wessex Formation (Barremian) of the
Isle of Wight. Journal of Vertebrate Paleontology. 29(2), 612-615.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed possible allosauroid (Crusafont-Pairo and Androver, 1966)
Late Tithonian-Middle Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (IPS-G1) maxillary tooth (82.7x33.4x16.9 mm)
Comments- This was originally referred to Carcharodontosaurus
(Crusafont-Pairo and Androver, 1966) and Megalosaurus (Kuhne and Crusafont-Pairo,
1968) before being described as a non-carcharodontosaurid allosauroid by Canudo
et al. (2006). Gasco et al. (2012) noted this tooth is very similar to "Megalosaurus"
ingens except in lacking enamel wrinkles.
References- Crusafont-Pairo and Androver, 1966. El primer representante
de la clase mamiferos hallado en el Mesozoico de Espana. Teruel. 35, 139-143.
Kuhne and Crusafont-Pairo, 1968. Mamiferos del Wealdiense de Una, cerca de Cuenca.
Nota preliminar. Acta Geologica Hispanica. 3(5), 133-134.
Canudo, Ruiz-Omenaca, Aurell, Barco and Cuenca-Bescos, 2006. A megatheropod
tooth from the Late Tithonian - Middle Berriasian (Jurassic-Cretaceous transition)
of Galve (Aragon, NE Spain). Neues Jahrbuch f�r Geologie und Pal�ontologie,
Abhandlungen. 239(1), 77-99.
Gasc�, Cobos, Royo-Torres, Mampel and Alcal�, 2012. Theropod teeth
diversity from the Villar del Arzobispo Formation (Tithonian-Berriasian) at
Riodeva (Teruel, Spain). Palaeobiodiversity and Palaeoenvironments. 92(2), 273-285.
unnamed possible allosauroid (Ruiz-Omenaca, Canudo and Infante, 2005)
Early Barremian, Early Cretaceous
Camarillas Formation, Spain
Material- (MPZ2005/316-317) teeth
Reference- Ruiz-Omenaca, Canudo and Infante, 2005. Presencia de un posible
Alosaurido (Dinosauria: Theropoda) en el Cretacico inferior (Barremiense Inferior)
de la Maca 3, (Galve, eruel). XXI Jornadas de la Sociedad Espanola de Paleontologia.
117-118.
unnamed possible allosauroid (Infante, Canudo, and Ruiz-Omenaca, 2005)
Early Barremian, Early Cretaceous
Mirambel Formation, Spain
Material- (LAD4r-1) tooth (~22 mm)
Reference- Infante, Canudo, and Ruiz-Ome�aca, 2005. First evidence
of theropod dinosaurs in the Mirambel Formation (lower Barremian, Lower Cretaceous)
in Castellote, Teruel. Geogaceta. 38, 31-34.
undescribed Allosauroidea (Gasulla, Ortega, Escaso and Sanz, 2006)
Late Barremian, Early Cretaceous
ANA site, Arcillas de Morella Formation, Spain
Material- (ANA37) tooth (?x12x6.3 mm) (Su�er and Santos-Cubedo, 2008)
Late Barremian, Early Cretaceous
Mas de la Parreta,
Arcillas de Morella Formation, Spain
(CMP-3-744) distal tibia (Gasulla, Ortega, Escaso and Sanz, 2006)
References- Gasulla, Ortega, Escaso and Sanz, 2006. Diversidad de ter�podos
del Cret�cico Inferior (Fm. Arcillas de Morella, Aptiense) en los yacimientos
del Mas de la Parreta (Morella, Castell�n). In Fern�ndez-Mart�nez
(ed.). XXII Jornadas de Paleontolog�a de la Sociedad Espa�ola
de Paleontolog�a. Libro de res�menes, 124-125.
Su�er and Santos-Cubedo, 2008. Dos dientes de ter�podo del yacimiento
ANA, Formaci�n Arcillas de Morella (Aptiense, Cret�cico Inferior,
Cinctorres, Castell�n). Encontro de Jovens Investigadores em
Paleontologia IV. Studia Geologica Salmanticensia. 8, 27-39.
unnamed allosauroid (Knoll, Buffetaut and Bulow, 1999)
Callovian, Middle Jurassic
Marnes de Dives, France
Material- (Bulow coll. 25192) braincase
.... frontals (Buffetaut and Enos, 1992)
Comments- Allain (2002) states this is probably an allosaurid. It is
not Piveteausaurus or Eustreptospondylus.
Reference- Buffetaut and Enos, 1992. Un nouveau fragment cr�nien
de dinosaure th�ropode du Jurassique des Vaches Noires (Normandie, France):
remarques sur la diversit� des th�ropodes jurassiques europ�ens
[A new theropod dinosaur skull fragment from the Jurassic of the Vaches Noires
(Normandy, France): Remarks on the diversity of the Jurassic theropods of Europe].
Comptes Rendus de l'Academie des Sciences Paris, S�rie II. 314, 217-222.
Knoll, Buffetaut and Bulow, 1999. A theropod braincase from the Jurassic of
the Vaches Noires Cliffs (Normandy, France): Osteology and palaeoneurology.
Bulletin de la Soci�t� g�ologique de France. 170(1), 103-109.
Allain, 2001. Redescription of Streptospondylus altdorfensis, Cuvier’s
theropod dinosaur, from the Jurassic of Normandy, Geodiversitas. 23(3), 349-367.
undescribed allosauroid (Allain, Vullo, Leprince, Neraudeau and Tournepiche,
2011)
Hauterivian-Barremian, Early Cretaceous
Angeac-Charente lignite, France
Material- (ANG 10-51) tooth (Neraudeau et al., 2012)
(ANG 10-262) tooth (Neraudeau et al., 2012)
Comments- Neraudeau et al. (2012) state these are indistinguishable from
Neovenator.
References- Allain, Vullo, Leprince, Neraudeau and Tournepiche, 2011.
An ornithomimosaur-dominated bonebed from the Early Cretaceous of Southwestern
France. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 61.
N�raudeau, Allain, Ball�vre, Batten, Buffetaut, Colin, Dabard,
Daviero-Gomez, Albani, Gomez, Grosheny, Le Loeuff, Leprince, Mart�n-Closas,
Masure, Mazin, Philippe, Pouech, Tong, Tournepiche and Vullo, 2012. The Hauterivian-Barremian
lignitic bone bed of Angeac (Charente, south-west France): Stratigraphical,
palaeobiological and palaeogeographical implications. Cretaceous Research. 37,
1-14.
undescribed possible allosauroid (Rich, Roland, Gangloff and Hammer,
1997)
Late Jurassic-Early Cretaceous
Suntar Series, Russia
Reference- Rich, Roland, Gangloff and Hammer, 1997. Polar Dinosaurs.
In Currie and Padian (eds.). Encyclopedia of Dinosaurs. Academic Press, New
York. 562-573.
undescribed possible allosauroid (Zhao and Xu, 2008)
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Material- (IVPP coll.) humerus (283 mm), radius, ulna (237 mm), metacarpal
I, phalanx I-1, manual ungual I (130 mm)
Comments- Zhao and Xu report this specimen has well developed humeral
internal and outer tuberosities; well developed entepicondyle; greatly expanded
deltopectoral crest; ectocondyle more expanded than entocondyle; high ulnohumeral
ratio of 84%; ulna slightly curved; smooth surface on the top of the well developed
olecranon process; well developed biceps tubercle on the radius; sub-rhombic
distal end of radius; manual ungual I 46% of humeral length. They say preliminary
analysis suggests it may be carnosaurian though it shares some characters with
coelurosaurs. The basal tyrannosauroid Yutyrannus was later described
from this formation and is of similar size with a similar ungual/humeral ratio,
but lacks an entepicondyle, has a larger entocondyle than ectocondyle, and a
shorter ulna (~73% of humeral length) which is straight. Thus it seems unlikely
Zhao and Xu's specimen is an early Yutyrannus record.
Reference- Zhao and Xu, 2008. A new theropod from the Early Cretaceous
Yixian Formation of Western Liaoning, China. Journal of Vertebrate Paleontology.
28(3), 164A.
unnamed possible Allosauroidea (Serrano-Martinez, Ortega, Sciscio, Tent-Manclus,
Bandera and Knoll, 2015)
Bathonian-Oxfordian, Middle-Late Jurassic
Tiouraren Formation of the Irhazer Group, Niger
Material- (TP4-6) tooth (55.9x20x13.3 mm)
(TP4-7) tooth (23.8x13.4x6.9 mm)
Comments- These grouped with Allosaurus and Acrocanthosaurus,
but no metriacanthosaurids were included.
Reference- Serrano-Martinez, Ortega, Sciscio, Tent-Manclus, Bandera and
Knoll, 2015. New theropod remains from the Tiourar�n Formation (?Middle
Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans.
Proceedings of the Geologists' Association. 126(1), 107-118.
unnamed allosauroid (Brusatte and Sereno, 2008)
Aptian-Albian, Early Cretaceous
Elrhaz Formation, Niger
Material- (MNN GAD1; holotype of Kryptops palaios in part) (~6-7
m; adult) fragmentary anterior dorsal vertebra (~70 mm), two partial mid dorsal
vertebrae, two ribs (~500-600 mm), sacrum (?,?,?,110,110 mm), ilia (650 mm),
pubes (~620 mm), ischia (~580 mm)
Comments- This was described as part of the holotype of the abelisaur
Kryptops by Sereno and Brusatte (2008). The postcrania were in situ,
while the maxilla was loose on the surface 15 meters away. Furthermore, Carrano
et al. (2012) noted the postcrania are too primitive for an abelisaur, suggesting
they do not belong together. They believed a a subrectangular fenestra between
the fourth and fifth sacral neural spines as in Giganotosaurus and a
peg-and-socket ilioischial articulation indicated the postcrania were carcharodontosaurid,
probably the contemporaneous Eocarcharia. Cau (online, 2012) ran the
postcrania as a separate OTU and found them to clade with Siamotyrannus
in Metriacanthosaurinae. They are retained here as Allosauroidea incertae sedis.
References- Sereno and Brusatte, 2008. Basal abelisaurid and carcharodontosaurid
theropods from the Lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica
Polonica. 53(1), 15-46.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Cau, online 2012. http://theropoda.blogspot.com/2012/05/tetanurae-update-part-2.html
undescribed possible allosauroid (Flynn, Simpson, Razafimanastoa, Andriatompohavana
and Totovolohy, 1997)
Middle Jurassic
Madagascar
Material- teeth, two presacral vertebrae
Comments- These were listed as belonging to a "small ?allosauroid",
but not described.
Reference- Flynn, Simpson, Razafimanastoa, Andriatompohavana and Totovolohy,
1997. New Triassic and Jurassic vertebrates from Madagascar. Journal of Vertebrate
Paleontology. 17(3), 46A.
unnamed possible allosauroid (Ghevariya and Srikami, 1994)
Middle Jurassic
Patcham Formation, India
Material- caudal vertebrae
Reference- Ghevariya and Srikarni, 1994. Dinosaur fauna from Mesozoic
rocks of Western India. Ninth International Gondwana Symposium. 143-163
Metriacanthosauridae Paul, 1988 sensu
Carrano, Benson and Sampson, 2012
Definition- (Metriacanthosaurus parkeri <- Allosaurus fragilis,
Carcharodontosaurus saharicus, Passer domesticus) (Hendrickx, Hartman
and Mateus, 2015)
Other definition- (Metriacanthosaurus parkeri <- Allosaurus fragilis,
Carcharodontosaurus saharicus, Meghalosaurus bucklandii, Spinosaurus aegyptiacus) (Rauhut and Pol, 2019)
= Sinraptoridae Currie and Zhao, 1994
Definition- (Sinraptor dongi <- Allosaurus fragilis, Carcharodontosaurus
saharicus) (Holtz et al., 2004)
Other definitions- (Sinraptor dongi <- Allosaurus fragilis)
(modified from Padian and Hutchinson, 1997)
(Sinraptor dongi <- Allosaurus fragilis, Monolophosaurus
jiangi,
(Sinraptor dongi <- Allosaurus fragilis, Carcharodontosaurus
saharicus, Passer domesticus) (Brusatte and Sereno, 2008)
= Sinraptoridae sensu Padian and Hutchinson, 1997
Definition- (Sinraptor dongi <- Allosaurus fragilis) (modified)
= Sinraptoridae sensu Sereno, 1998
Definition- (Sinraptor dongi <- Allosaurus fragilis, Monolophosaurus
jiangi,
= Sinraptoridae sensu Holtz et al., 2004
Definition- (Sinraptor dongi <- Allosaurus fragilis, Carcharodontosaurus
saharicus)
= Sinraptoridae sensu Brusatte and Sereno, 2008
Definition- (Sinraptor dongi <- Allosaurus fragilis, Carcharodontosaurus
saharicus, Passer domesticus)
= Metriacanthosaurus sensu Rauhut and Pol, 2019
Definition-
Brusatte and Sereno's (2008) definition of Sinraptoridae differs from Holtz
et al.'s (2004) by including Passer as an external specifier, which I
view as superfluous, since a (Allosaurus, Carcharodontosaurus
(Sinraptor, Passer)) topology has never been advocated. Megalosaurus
might be a better choice for a tertiary external specifier, to cover traditional
phylogenies prior to 1993.
In several phylogenies prior to 1994, metriacanthosaurids such as Yangchuanosaurus
were seen as outside Avetheropoda. While not seen in many recent analyses, this
only takes five more steps when constrained in Carrano et al.'s (2012) matrix,
so is still quite possible.
Not metriacanthosaurids- Bakker et al. (1992) thought the maxilla OUMNH
J.13506 was a 'yangchuanosaur', but this was referred to Megalosaurus bucklandii
by Benson (2010).
References- Paul, 1988. Predatory Dinosaurs of the World. Simon &
Schuster, New York. 464 pp.
Bakker, Kralis, Siegwarth and Filla, 1992. Edmarka rex, a new, gigantic
theropod dinosaur from the Middle Morrison Formation, Late Jurassic of the Como
Bluff outcrop region. With comments on the evolution of the chest region and
shoulder in theropods and birds, and a discussion of the five cycles of origin
and extinction among giant dinosaurian predators. Hunteria. 2(9), 1-24.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Chienkosaurus Young,
1942
C. ceratosauroides Young, 1942
Tithonian?, Late Jurassic
IVPP locality 47, upper Guangyuan Group, Sichuan, China
Lectotype- (IVPP V237A) (~8 m) posterior premaxillary tooth (44x16x12 mm)
Referred- ?(IVPP V193) ulna (164 mm) (Young, 1942)
Bathonian-Callovian?, Middle Jurassic
IVPP locality 49, middle Guangyuan Group, Sichuan, China
?(IVPP V190) (~5 m) ~ninth caudal centrum (66 mm) (Young, 1942)
Other diagnoses- Young (1942) originally diagnosed Chienkosaurus
with- "Teeth thick and sharply pointed with fine palisade
denticulations on both sides. The anterior which are finer than the
posterior ones push lingually
towards the base and form a ridge topping at a distance before the base of the tooth."
Comments-
The material was discovered in late Spring 1941, with the type
consisting of four isolated teeth IVPP V237A-D. Young's (1942)
diagnosis was "Mainly based upon" the largest tooth (V237A), with the
three smaller teeth considered immature and (possibly incorrectly)
lacking their bases. He stated "The general shape of the teeth
resembles that of Labrosaurus stechowi" which was prescient as both are based on mesial dentition, and considered Chienkosaurus a ceratosaurid based on the questionably referred postcrania. Ironically, "Labrosaurus" stechowi is now thought to be ceratosaurid, but as Young noted Chienkosaurus lacks its lingual fluting which has proven to be a ceratosaurid character. Subsequently, Chienkosaurus
was generally placed in Megalosauridae (e.g. Romer, 1956; Steel, 1970;
Dong et al., 1978) when it was used as a waste basket for almost all
large Jurassic theropods including Ceratosaurus and later Yangchuanosaurus. Note Huene (1959) when citing Chienkosaurus as named in 1958 from the Late Cretaceous of Shantung meant to list Chingkankousaurus. Dong et al.
(1983) reported that "Rozhdestvensky (1964) proposed that the four
teeth of Chienkosaurus could
possibly belong to the Crocodilia" (translated), but which work this
corresponds to was not listed in the bibliography and cannot be determined. Dong et al.
also stated "during the editing of "The Handbook of Chinese Fossil
Vertebrates," Zhiming Dong conducted a review of these four specimens
and formally confirmed that the best preserved tooth among the V237
collection was a premaxillary tooth of a carnosaurian dinosaur, but
that the remaining three teeth were assignable to the crocodile Hsisosuchus." The dentition of Hsisosuchus
has not been described or figured in enough detail to distinguish it
from theropods, but two of the teeth (IVPP V237B and V237D)
are similar in being short and barely recurved with a high crown base
ratio, characters shared with the tooth figured separately in
Hsisosuchus' type description. They are provisionally placed in Hsisosuchus sp. here. The third supposed Hsisosuchus
tooth (IVPP V237C) is different in having a distinctly D-shaped section
with strong carinae somewhat like Guimarota tyrannosauroid premaxillary
tooth IPFUB GUI D 89, so is provisionally placed in Tyrannosauroidea
here. Retaining only one of Chienkosaurus'
syntype teeth in the genus would make it the lectotype, and as ICZN
Article 74.5 states "In a lectotype designation made before 2000,
either the term "lectotype", or an exact translation or equivalent
expression (e.g. "the type"), must have been used or the author must
have unambiguously selected a particular syntype to act as the unique
name-bearing type of the taxon", and Dong et al. explicitly make Chienkosaurus a synonym of Szechuanosaurus, and of Chienkosaurus'
syntypes only consider IVPP V237A to be theropodan, this is here
considered a valid lectotype designation. Rozhdestveksy (1977) earlier
listed Szechuanosaurus campi and Chienkosaurus ceratosauroides as "synonyms?" in his Table 1 without comment, while Dong et al.'s synonymization was based on examining Yangchuanosaurus
teeth from CV 00214 to correctly determine "the differences among
carnosaur dentitions are due only to being in a different position in
the dentition" and noting Chienkosaurus' and Szechuanosaurus'
types are from the same locality. While this indeed makes it possible
they even derive from the same individual, none of the teeth have been
shown to be diagnostic within metriacanthosaurids, and synonymization
should be based on autapomorphies or unique combinations of characters
instead of provenance. This synonymization of part of the Chienkosaurus type with Szechuanosaurus
was followed by Molnar et al. (1990) where they consider the taxon an
allosaurid, which makes sense as Dong was a coauthor. Most recently,
Hendrickx et al. included Chienkosaurus
in their cluster analyses, although the taxon is never mentioned in the
text, matrices or table of examined taxa. Classical/Hierarchical
clustering resolves it with Genyodectes, Sinraptor dongi (the only metriacanthosaurid analyzed there) and Allosaurus, while neighbour joining clustering resolves it sister to a clade whose basal members are 'Indosuchus' AMNH jaws, Allosaurus and S. dongi.
Young placed locality 47 at "the top part of the Kuangyuan Series and
immediately below the Chentsianyen conglomerate", now known as the
Guangyuan Group and the Chengqiangyan Group, with the former
corresponding to the Xiashaximiao Formation through the Penglaizhen
Formation. As it was found "immediately below" the boundary (layer 8b in Young et al., 1943), Chienkosaurus
may be from the Penglaizhen Formation or slightly lower Shuining
Formation. The age is listed as Tithonian on fossilworks and in
Weishampel (1990), the latter cited as from "Dong (pers. comm.)".
Geographically, locality 47 is "the hill slopes under the so-called
Chentsianyen escarpment S. of the Kuangyuan city", now known as
Guangyuan.
The tooth is similar to many large theropod teeth in general
characters, but is from the premaxilla as evidenced by the twisted
mesial carina and reduced extent of mesial serrations. The crown
base ratio (.75) is between the third and fourth premaxillary teeth of
Sinraptor dongi's holotype
(pm3 .60; pm4 1.04), which also match in size (pm3 FABL 16.87 mm; pm4
BW 12.26 mm) and in lacking mesial serrations basally. The cited
mesial (15 per 5 mm) and distal (6.7-10 per 5 mm) serration densities
are matched by teeth of S. dongi,
and the strong mesial carina Young describes could easily be due to the
"longitudinal groove adjacent to the mesial carina, on the lingual
surface of the crown" "clearly present in lpm3 and lpm4" as described
by Hendrickx et al. (2020) for S. dongi. Thus Chienkosaurus is indistinguishable from Sinraptor dongi
as far as can be determined from the description, and given its poorly
constrained age could be contemporaneous or even synonymous. Hendrickx
et al.'s matrices show no differences between Yangchuanosaurus shangyouensis (including Y. magnus), Sinraptor dongi and S. hepingensis that can be evaluated for Chienkosaurus,
so pending Hendrickx's in prep. study on metriacanthosaurid dental
anatomy the genus is considered Metriacanthosauridae indet..
Referred material- Young (1942) figured and described an ulna from the type locality (IVPP V193), stating he "would prefer to refer this ulna to Chienkosaurus ceratosauroides above described." The element is very different from Limusaurus
in having marked transverse expansions proximally and distally as well
as a triangular versus reniform proximal end, so that if Sinocoelurus is closely related to that genus the ulna is unlikely to belong to it. Eoabelisaurus has a much longer olecranon. The ulnae of megalosaurids and Kaijiangosaurus are far more robust with more proximally extended olecranons, while those of most coelurosaurs (e.g. Zuolong, Guanlong, Coelurus, Tanycolagreus, Fukuivenator) are much more slender with developed olecranons as well. Fukuiraptor
has a dissimilar ulna with a strong olecranon, prominent anteroproximal
longitudinal ridge and unexpanded distal end. This leaves several
roughly comparable taxa whose ulnae have been figured in
anteroposterior view- Ceratosaurus, Poekilopleuron, Yangchuanosaurus, Allosaurus and Haplocheirus. Young compared it favorably to the former, writing "it fits rather well with the ulna of Ceratosaurus nasicornis
(length of ulna, 17.7 cm.) which is only slightly longer than the
present form", and indeed the main difference in profile is the more
gradual proximal expansion laterally. However, in proximal view IVPP
V193 differs from Ceratosaurus and most other proximally figured ulnae in having a centrally placed olecranon (also seen in Coelurus, but not Tanycolagreus). While only photographed in anterior view, the ulna of Yangchuanosaurus (CV 00214) would also seem to have a centrally placed olecranon, so IVPP V193 may be correctly referred to Chienkosaurus/Szechuanosaurus after all.
Young
(1942) describes IVPP V190 as "A complete centrum of an anterior caudal
vertebra (or posterior lumbar) with length 66 mm., breadth 41 mm.,
minimum breadth of the centrum 24 mm", noting it "fit in size with Chienkosaurus ceratosauroides" and calling it Theropoda indet. in the plate caption but also saying there it "probably belonging to Chienkosaurus ceratosauroides." With a length/height ratio of 144% it is comparable to the ninth caudal of Sinraptor hepingensis
and indistinguishable in lateral view. It differs in being 85% wider
than tall vs. 95%, but this is within the range of variation in hepingensis'
caudals. Notably, this is from a different locality than the
type,
said by Young to be in "the middle part of the" ... "Kuangyuan Series",
layer 5a in Young et al. (1943),
and thus possibly corresponding to the Shangshaximiao Formation.
IVPP locality 49 is described as being "a few kilometers N. of the city
[Guangyuan] in the hills along the Chengtu-Sian highway", and the
highway connecting Chengdu and Xi'an (as the cities are now called) is
China National Highway 108, or G108. Thus
while lacking a plausible connection to Chienkosaurus, it is congruent with being metriacanthosaurid but may also be e.g. piatnitzkysaurid or megalosaurid.
References- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan,
China. Bulletin of the Geological Society of China. 22(3-4), 293-309.
Young, Bien and Mi, 1943. Some geologic problems of the Tsinling. Bulletin
of the Geological Society of China. 23(1-2), 15-34.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 1-772.
Huene, 1959. Saurians in China and their relations. Vertebrata PalAsiatica. 3(3), 119-123.
Steel, 1970. Part 14. Saurischia. Encyclopedia of Paleoherpetology. Gustav Fischer Verlag. 1-87.
Rozhdestvensky, 1977. The study of dinosaurs in Asia. Journal of the Palaeontological Society of India. 20, 102-119.
Dong, Zhang, Li and Zhou, 1978. [A new carnosaur discovered in
Yongchuan, Sichuan]. Chinese Science Bulletin. 23(5), 302-304.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 169-209.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.
Hendrickx, Stiegler, Currie, Han, Xu, Choiniere and Wu, 2020. Dental anatomy of the apex predator Sinraptor dongi (Theropoda: Allosauroidea) from the Late Jurassic of China. Canadian Journal of Earth Sciences. 57(9), 1127-1147.
"Yuanmouraptor" Anonymous, 2014
Middle Jurassic
Yuanmou County, Yunnan, China
Material- (ZLJ 0115) partial skull, mandibles (one incomplete, one partial),
postcrania
Comments- This specimen is on display at the ZLJ as a new carnosaur,
but has yet to be described. There is a mounted skeleton, but how much is original
is unreported. Hendrickx et al. (2019) call this "an
undescribed metriacanthosaurid (ZLJT 0115)", state it has mesial and
lateral teeth with "four to six, possibly more" flutes, and list it as
"Metriacanthosauridae indet." in their Appendix 1 indicating
information was from photos provided by Stiegler.
References- Anonymous, 2014. Special Exhibition: Legends of the Giant
Dinosaurs. Hong Kong Science Museum newsletter. 1-3-2014, 2-7.
Hendrickx, Mateus, Ara�jo and Choiniere, 2019. The distribution of
dental features in non-avian theropod dinosaurs: Taxonomic potential,
degree of homoplasy, and major evolutionary trends. Palaeontologia
Electronica. 22.3.74, 1-110.
undescribed Metriacanthosauridae (Gerke and Wings, 2014)
Kimmeridgian, Late Jurassic
Langenberg Quarry and/or Hannover, Germany
Material- (NLMH coll.) teeth
Reference- Gerke and Wings, 2014. Characters versus morphometrics: A
case study with isolated theropod teeth from the Late Jurassic of Lower Saxony,
Germany, reveals an astonishing diversity of theropod taxa. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 137.
unnamed possible metriacanthosaurid (Young and Sun, 1957)
Kimmeridgian-Tithonian, Late Jurassic
Langgou IVPP 93008-2?, Kalaza Formation, Xinjiang, China
Material- (IVPP V903) (~10 m) (mandible ~1.17 m) anterior dentary
Diagnosis- Provisionally indeterminate compared to Sinraptor? hepingensis.
Comments-
The specimen was discovered in September 1955 (Chinese text of Young
and Sun, 1957). Young and Sun described IVPP V903 as being from
"Ying-choe-shih some 13 kilometers S. E. of Pichan, east of the Turfan
basin, Sinkiang", but Pichan is now known as Shanshan County while
Sinkiang is Xinjiang. Dong (1977) lists this locality as
"Yingzuishigou (Langgou) southwest of Qiketai, Shanshan District"
(translated), which can be correlated with the type locality of
Hudiesaurus, given by Dong (1997b) as "Langgou (wolf groove), Qiketia
area, Shanshan County of the Turpan Basin." Dong (1997a) indicates
this is IVPP locality 93008-2, but Dong (1992) stated "The beds
yielding the fossil were unknown; the fossil was collected by local
people", so a more precise locality is unlikely to become known.
Similarly, while Young and Sun initially reported the beds as
Cretaceous, Dong (1977) first notes that "the time of the
Szechuanosaurus campi Young should be of the Later Jurassic Hungshan
series, not of the early Cretaceous Tugulu series. Wei Jingmin also
verifies this point in his recent letter." Indeed, Dong (1987) reports
it is from "the late Jurassic Kelaza Formation", currently called the
Kalaza Formation. Dong (1992) incorrectly calls it "the middle part of
a left maxilla", which would make no sense as the shorter alveolar wall
would be lateral and the dorsal margin would be rounded in section, the
latter showing this isn't even the anterior part of a right maxilla.
Upchurch et al. (2021) repeat this mistake when referring to "part of a
left maxilla of a ‘megalosaurid’" from Qiketai. Dong (1987)
photographs the medial surface in color on page 70 as "the lower jaw of
cf. Szechuanosaurus campi",
which shows slight breakage has occured since 1957 and that the
Meckelian groove is deeper than suggested by Young and Sun's plate.
Based on dentary depth (~88 mm) we might expect it to be ~31% larger
than Sinraptor dongi's holotype, although dentary depth seems likely to
increase allometrically with size in large theropods, so perhaps it was
a bit shorter but more robust.
Young and Sun (1957) only compared it to Chinese large theropods known at the time (Szechuanosaurus syntypes and referred teeth from Laiyang in the Wangshi Group, Chienkosaurus and Bohlin's Inner Mongolia ?Prodeinodon),
concluding "Comparing specimen by specimen of all those mentioned
Chinese forms kept in our laboratory the Sinkiang specimen is
doubtlessly closer to the Szechuanosaurus. We thus refer our specimen as cf. Szechuanosaurus campi."
Earlier they stated that "The serration is rather densely arranged,
about 18-26 per centimeter which is much less than the type of Szechuanosaurus and rather close to that of the teeth of Laiyang referred to Szechuanosaurus", which are far too late to belong to the genus. Not only this, Chienkosaurus was distinguished from Szechuanosaurus (and thus IVPP V903?) in part by its "smaller size, straightness and bluntness" which characterize the ?Hsisosuchus and ?tyrannosauroid teeth (IVPP V237B-D) no longer placed in the taxon, while "Prodeinodon mongoliense
[sic] and other carnivorous dinosaur from Inner Mongolia are too scant
for a precise comparison", so even a comparison to Laiyang "Szechuanosaurus"
teeth was based on very little. Comparison to Szechuanosaurus is
limited as detailed descriptions of any of these teeth are lacking, but
the serration density (9-13 per 5 mm) is similar to at least
Szechuanosaurus syntypes IVPP V235 (8.5 per 5 mm) and V236 (12 per 5
mm), and the replacement tooth of alveolus 4 (III4 in Young and Sun's
figure) is similar in side outline to IVPP V238B. On the other hand,
IVPP V235 and V239 are more recurved, and the latter more acute, than
any preserved IVPP V903 teeth, and V238A and V239 have finer serrations
(17-~19 per 5 mm). The size is similar, with the most complete tooth
(II1 of Young and Sun) having a FABL of ~21 mm just like IVPP V236, and
the geological ages of the Kalaza Formation and upper Guangyuan Group
may be comparable. Thus IVPP V903 may belong to the same taxon as at
least some of the Szechuanosaurus campi syntypes (IVPP V236, 238B), but
side outline and serration density (not even specified to carina, let
alone basoapical position) will rarely be sufficient to refer a tooth
to a genus of large theropod. Note a tooth from Nanhuxiang in the Kalaza Formation (IVPP coll.) figured by Dong (1977) as Szechuanosaurus campi resembles S. campi
syntype IVPP V238A in side outline and based on provenence may belong
to the same taxon as IVPP V903, but in the absense of more data is
placed in Averostra indet. here.
Young and Sun (1957) proposed two potentially characteristic features
of IVPP V903. First, "the dental foramens below the tooth row are not
arranged in straight line, but are jumping anteriorly and upwardly in
pair almost regularly", which is also seen in widespread taxa such as
ceratosaurids, Duriavenator, Dubreuillosaurus, Sinraptor dongi,
carcharodontosaurines, Yutyrannus, Nuthetes, Dromaeosaurus and Deinonychus. Second,
"the jaw is obviously thickening at the middle as seen clearly in the
cross-section", referencing the 'lateral shelf' later described by
Molnar (1974). While the degree of its development is impossible to
derive from figures without a coronal section, other taxa with a
dorsally angled surface at the level of the primary neurovascular
foramen line include not only Labocania, but carcharodontosaurines,
metriacanthosaurids, Kelmayisaurus, Magnosaurus, and perhaps
Marshosaurus and Jinbeisaurus among large theropods. While
therizinosauroids have dentary shelves (as noted by Chure, 2000),
Falcarius shows these evolved after phyllodont teeth, and the shelf
structure in e.g. Erlikosaurus is quite different from IVPP V903 and
the taxa listed above, being at or above the alveolar edge and not even
developing until after the fifth tooth, which is more posteriorly
placed due to the edentulous symphysis. The less angled structure in
IVPP V903 is thus referred to as a dentary step below.
The next author to comment on IVPP V903 specifically was Molnar (1974),
who in his description of the new theropod Labocania stated "Such a
lateral [dentary] shelf is not unkown [sic] in theropods, having been
described for at least one other, cf. Szechuanosaurus campi from
Sinkiang" and correctly notes later that "It is also not clear, and was
not clear to the describers, Young and Sun, that the dentary fragment
from Sinkiang actually pertains to Szechuanosaurus." The only direct
comparison given is that "It does appear that the dentary fragment of
cf. S. campi was about 5 cm in maximal width (Young and Sun, 1957, fig.
2), which is pretty close to that (6 cm) of the dentary fragment of
Labocania" (scaling figures today, IVPP V903 is ~48 mm while Labocania
has a broken lateral edge so is 65+ mm). However, the dentary fragment
of Labocania is from the posterior portion and thus not comparable to
IVPP V903 which only covers the first four alveoli, so that differences
in sectional shape from the latter (more prominent overhang of
Meckelian groove/fossa; vertically concave medial surface above this)
could merely be due to position. The teeth in both specimens are only
briefly described but have similar serration densities (9-13 per 5 mm
in IVPP V903 vs. 10-14 mesially and 11-14 distally in Labocania).
Most recently, Chure (2000) agreed "there is no justification for
referring this specimen to Szechuanosaurus", and suggested the lateral
step and labial foramina placed "close to the dorsal margin of the
dentary" are unusual features. The latter seems to be exaggerated by
dorsal breakage and is only true of the first four foramina, which is
also found in Ceratosaurus, Monolophosaurus, Piatnitzkysaurus,
Dubreuillosaurus, all(?) carnosaurs including Sinraptor, Allosaurus and
carcharodontosaurids, Fukuiraptor, Nuthetes and Eotyrannus. Despite
these features, Chure considered IVPP V903 to be "Theropoda
indeterminate" and it has never been compared in detail to other
theropods.
Comparisons- IVPP V903 can be
excluded from clades with derived anterior dentary morphologies like
noasaurids, spinosaurids and maniraptoromorphs besides eudromaeosaurs,
with most maniraptoromorphs also being far too small with rare
exceptions. Compared to ceratosaurids, IVPP V903 differs in having a
lateral step and from Ceratosaurus itself in having no lingual grooves
on dentary tooth 2, and Genyodectes further differs in having a chin under tooth 3. Monolophosaurus differs in lacking the lateral
step, having labial foramina arranged in an arc, a Meckelian groove
that ends under tooth 3, and a higher lingual wall. Marshosaurus is
quite distinct in having an anterodorsally rounded and dorsoventrally
expanded dentary with fewer and larger foramina more ventrally
positioned anteriorly. Piatnitzkysaurus lacks a lateral step, has two
rows of labial foramina and a Meckelian groove that ends under tooth
3. Eustreptospondylus lacks a lateral step, has a convex ventral edge
under alveoli 2-4, and has a more ventrally placed Meckelian groove
that ends under tooth 4. Duriavenator and Megalosaurus lack a lateral
step, have randomly arranged labial foramina, and a much lower
anteroventrally angled Meckelian groove that ends under tooth
3, and Megalosaurus further differs in having a chin under tooth 3. Torvosaurus lacks a lateral
step, has a convex ventral edge under alveoli 2-4, and has an extremely
shallow Meckelian groove that isn't obvious as far anterior as alveolus 7. Dubreuillosaurus lacks a lateral step, has a
convex ventral edge under alveoli 2-4, has two rows of labial foramina,
a higher lingual wall and a much lower anteroventrally angled Meckelian
groove. Magnosaurus has a convex ventral edge under alveoli 2-4, a
lower lingual wall and a much lower anteroventrally angled
Meckelian groove. Australovenator lacks a lateral step, and it and
Fukuiraptor have a much lower Meckelian groove. Proceratosaurids and
Dilong are difficult to compare being articulated with the skull, but both lack lateral steps at least.
Nuthetes lacks a lateral step and has a much lower anteroventrally
angled
Meckelian groove. Eotyrannus lacks a lateral step, has an anterodorsal
alveolar notch, an extremely shallow Meckelian groove that ends under tooth 4 and a lower
lingual wall. Jinbeisaurus has smaller anterior teeth so that three
alveoli are anterior to the chin and a Meckelian groove that ends under tooth 4. Dromaeosaurids (represented by
Deinonychus and Dromaeosaurus) lack a lateral step and have a much lower
Meckelian groove. Asfaltovenator lacks a lateral step, and has a
convex ventral edge under alveoli 2-4. Kelmayisaurus has a convex
ventral edge under alveoli 2-4, an accessory labial groove at
midheight, and a lower anteroventrally angled
Meckelian groove. Allosaurus lacks a lateral step, has a convex
ventral edge
under alveoli 2-4, and a lower
Meckelian groove. Neovenator lacks a lateral step, has two rows of
labial foramina, and a lower Meckelian groove. Acrocanthosaurus has
the labial foramina in a groove, and a lower Meckelian groove that ends under tooth
3.
Carcharodontosaurines differ in having anteriorly diverging Meckelian
and paradental grooves, while Carcharodontosaurus iguidensis itself has
a lower lingual wall, lower Meckelian groove and second row of labial
foramina; and Giganotosaurus and Tyrannotitan both have lower
anteroventrally angled
Meckelian grooves but Mapusaurus does not. Yangchuanosaurus is an
important taxon to compare IVPP V903 with given the standard dating of
the Shangshaximiao Formation and geographical proximity, but any
comparisons are limited by the type's dentary being partially hidden by
the articulated upper jaw and CV 00216's anterior dentary only being
figured somewhat schematically in medial view. Regardless, it lacks a
lateral step until perhaps the posterior half of the alveolar margin
and has a convex ventral edge under alveoli 2-4, but in medial view the
angle, height and anterior extent of the Meckelian groove are similar as is the height
of the lingual wall. Sinraptor dongi is almost identical except the
portion anterior to the chin is more elongate and the groove ventral to
the Meckelian groove is deeper. S. hepingensis is more similar in
having a shorter pre-chin section and lacking a groove below the
Meckelian groove, but the anterior extent of its lateral step is less
obvious.
Thus of theropods with laterally and medially figured anterior
dentaries, IVPP V903 is most similar to Sinraptor and
Mapusaurus. S. dongi has similar serration densities in mesial dentary
teeth (10-18 per 5 mm; average 11-14.6 vs. 9-13 in IVPP V903), as does
Mapusaurus in isolated teeth (7-15 per 5 mm).
Importantly, Mapusaurus is Late Cretaceous and
other
carcharodontosaurids are less similar. Metriacanthosaurids are very common in
Middle-Late Jurassic China on the other hand, making them the most
likely identity for IVPP V903. While seemingly more similar to Sinraptor than Yangchuanosaurus, it may be more recent than named species of either and contemporaneous/synonymous with more poorly resolved taxa like Chienkosaurus and Yangchuanosaurus? "longqiaoensis" instead. Because of this and the poor existing descriptions of Yangchuanosaurus dentaries, IVPP V903 is merely placed as Metriacanthosauridae here.
References- Young and Sun, 1957. Note on a fragmentary carnosaurian mandible
from Turfan, Sinkiang. Vertebrata PalAsiatica. 1(2), 2027-2036.
Molnar, 1974. A distinctive theropod dinosaur from the Upper Cretaceous of Baja
California (Mexico). Journal of Paleontology. 48(5), 1009-1017.
Dong, 1977. On the dinosaurian remains from Turpan, Xinjiang.
Vertebrata PalAsiatica. 15(1), 59-66.
Dong, 1987. Dinosaurs from China. China Ocean Press. 114 pp.
Dong, 1992. Dinosaurian Faunas of China. Ocean Press/Springer-Verlag.
188 pp.
Dong, 1997a. Vertebrates of the Turpan Basin, the Xinjiang Uygur
Autonomous Region, China. The Geology of the Turpan Basin. In Dong
(ed.). Sino-Japanese Silk Road Dinosaur Expedition. China Ocean Press.
96-101.
Dong, 1997b. A gigantic sauropod (Hudiesaurus sinojapanorum,
gen. et sp. nov.) from the Turpan Basin, China. In Dong (ed.).
Sino-Japanese Silk Road Dinosaur Expedition. China Ocean Press. 102-110.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Upchurch, Mannion, Xu and Barrett, 2021. Re-assessment of the Late Jurassic eusauropod dinosaur Hudiesaurus sinojapanorum
Dong, 1997, from the Turpan Basin, China, and the evolution of
hyper-robust antebrachia in sauropods. Journal of Vertebrate
Paleontology. 41(4), e1994414.
undescribed metriacanthosaurid (Chanthasit and Suteethorn, 2013)
?Oxfordian-Early Valanginian, ?Late Jurassic-Early Cretaceous
Phui Noi, Phu Kradung Formation, Thailand
Material- (Phui Noi metriacanthosaurid) (SM-KS34-1498) maxilla (520 mm)
....(SM-KS34 coll.) (at least 2 individuals) three premaxillae, three
maxillae, teeth, ulna, metatarsal II, two metatarsals III, pedal
phalanges
Comments- Chanthasit and
Suteethorn (2013) mention a maxilla with "15 maxillary alveoli", and
say "the lateral surface of the maxilla is rugose only along its
anterior edge and just above the tooth row" and that "the upper part and the border of the antorbital
fossa are smooth and indented" with "a complex of accessory openings,
which is characteristic of the Sinraptoridae." It's photographed
in
Samathi et al. (2019). They also mention "metatarsal II,
metatarsal
III, and phalanges referred to the right pes of medium to large-sized
theropod." Samathi et al. (2016) suggest it "appears to have a
robust
and a gracile morph, possibly due to sexual dimorphism." Samathi
et al. (2019) state "the cranial material is currently under study."
Samathi (2016) wrote "preliminary analysis suggested the Phu Noi
specimens are closer to Sinraptor dongi ... than Yangchuanosaurus." On
the other hand, Samathi et al. (2016) use Carrano et al.'s tetanurine
matrix to determine "the Phu Noi specimen to represent a new taxon
nested within Metriacanthosauridae. The Phu Noi metriacanthosaurid may
not be a metriacanthosaurine since the anteroventral border of the
maxillary antorbital fossa is not demarcated by a raised ridge."
References- Chanthasit and
Suteethorn, 2013. The first nearly complete upper jaw (maxilla) of a
large-sized theropod dinosaur from Phu Kradung Formation (Late
Jurassic-Early Cretaceous) of Thailand. The 3rd International
Conference on the Palaeontology of Southeast Asia. Abstract, 22.
Samathi, 2016. Metriacanthosaurids (Dinosauria: Theropoda) from
Thailand and paleobiogeography of Metriacanthosauridae. XIV EAVP
Meeting. 213.
Samathi, Chanthasit and Sander, 2016. New material of a new
metriacanthosaurid (Dinosauria, Theropoda) from the Phu Noi locality
(Late Jurassic-Early Cretaceous) of Thailand. Journal of Vertebrate
Paleontology. Program and Abstracts, 217.
Samathi, Chanthasit and Sander, 2019. A review of theropod dinosaurs
from the Late Jurassic to mid-Cretaceous of southeast Asia. Annales de
Pal�ontologie. 105(3), 201-215.
Yangchuanosaurus
Dong, Zhang, Li and Zhou, 1978
= "Szechuanoraptor" Chure, 2000
Comments- Rauhut (2000) believed Yangchuanosaurus and Sinraptor
to be "almost identical" and found it likely all specimens were from
a single species, which would be Yangchuanosaurus shangyouensis. More
recently, Carrano et al. (2012) found Sinraptor to be closer to Metriacanthosaurus,
Siamotyrannus and Shidaisaurus than to Yangchuanosaurus.
References-
Dong, Zhang, Li and Zhou, 1978. [A new carnosaur discovered in
Yongchuan, Sichuan]. Chinese Science Bulletin. 23(5), 302-304.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (UT-CO) and a revision of the theropod family
Allosauridae. PhD thesis. Columbia University. 964 pp.
Rauhut, 2000. The interrelationships and evolution of basal
theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Y. shangyouensis Dong, Zhang, Li and Zhou, 1978
= Szechuanosaurus "yandonensis" Dong, Zhang, Li and Zhou, 1978
= Yangchuanosaurus magnus Dong, Zhou and Zhang, 1983
= Metriacanthosaurus "carpenteri" Paul, 1988
= Metriacanthasaurus shangyouensis (Dong, Zhang, Li and Zhou, 1978) Paul,
1988
= “Szechuanoraptor dongi” Chure, 2000
Bathonian-Callovian, Middle Jurassic
Shangyou Reservoir, Shangshaximiao Formation, Sichuan, China
Holotype- (CV 00215) (7.9 m, 1.33 tons, subadult) skull (780 mm), mandibles
(780 mm), axis (64 mm), third cervical vertebra (78 mm), fourth cervical vertebra
(95 mm), fifth cervical vertebra (115 mm), sixth cervical vertebra 118 mm),
seventh cervical vertebra (120 mm) eighth cervical vertebra (138 mm), ninth
cervical vertebra (114 mm), tenth cervical vertebra (95 mm), fourteen cervical
ribs (eighth 500 mm), first dorsal vertebra (120 mm), second dorsal vertebra
(120 mm), third dorsal vertebra (120 mm), fourth dorsal vertebra (120 mm), fifth
dorsal vertebra (125 mm), sixth dorsal vertebra (130 mm), seventh dorsal vertebra
(130 mm), eighth dorsal vertebra (128 mm), ninth dorsal vertebra (130 mm), tenth
dorsal vertebra (130 mm), eleventh dorsal vertebra (135 mm), twelfth dorsal
vertebra (132 mm), thirteenth dorsal vertebra (133 mm), twenty-four dorsal ribs
(100-1080 mm), first sacral vertebra (130 mm), second sacral vertebra (110 mm),
third sacral vertebra (91 mm), fourth sacral vertebra (100 mm), fifth sacral
vertebra (105 mm), first caudal vertebra (98 mm), second caudal vertebra (102
mm), third caudal vertebra (100 mm), fourth caudal vertebra (96 mm), fifth caudal
vertebra (106 mm), sixth caudal vertebra (118 mm), seventh caudal vertebra (108
mm), eighth caudal vertebra (107 mm), ninth caudal vertebra (115 mm), tenth
caudal vertebra (110 mm), eleventh caudal vertebra (110 mm), twelfth caudal
vertebra (110 mm), twelve chevrons (to 199 mm), distal scapula, fragmentary
humerus, ilia, pubes, ischia, femora (850 mm), tibiae (755 mm), fibulae, astragalus,
calcaneum, several pedal phalanges(?)
Bathonian-Callovian, Middle Jurassic
Hongjiang Machine Factory, Shangshaximiao Formation, Chongqing, China
Referred- (CV 00216; holotype of Yangchuanosaurus magnus) (10.5 m; 3.1 tons) partial
skull (1.11 m), mandibles (1.17 m), axis (115 mm), third cervical vertebra,
fifth cervical vertebra, sixth cervical vertebra, eighth cervical vertebra,
tenth cervical vertebra, first dorsal vertebrae, second dorsal vertebra, sixth
dorsal vertebra, eighth dorsal vertebra, tenth dorsal vertebra, twelfth dorsal
vertebra, first sacral vertebra (145 mm), second sacral vertebra (135 mm), third
sacral vertebra (120 mm), fourth sacral vertebra (111 mm), fifth sacral vertebra
(120 mm), first caudal vertebra (130 mm), second caudal vertebra (125 mm), third
caudal vertebra (125 mm), fourth caudal vertebra (120 mm), four mid caudal vertebrae,
ilium, incomplete ischium, femur (950 mm), pedal phalanx I-1, pedal phalanx
III-1
Bathonian-Callovian, Middle Jurassic
Wujiaba, Shangshaximiao Formation, Sichuan, China
(CV 00214; intended holotype of "Szechuanoraptor dongi"; Metriacanthosaurus "carpenteri"; Szechuanosaurus "yandonensis") (3.8 m, 130
kg) many anterior and lateral teeth, axis (55 mm), third cervical vertebra (60
mm), fourth cervical vertebra (65 mm), fifth cervical vertebra (55 mm), sixth
cervical vertebra (65 mm), seventh cervical vertebra (70 mm), eighth cervical
vertebra (62 mm), first dorsal vertebra (60 mm), second dorsal vertebra (60
mm), third dorsal vertebra (74 mm), fourth dorsal vertebra (60 mm), seventh
dorsal vertebra (65 mm), eighth dorsal vertebra (62 mm), ninth dorsal vertebra
(62 mm), tenth dorsal vertebra, dorsal ribs, gastralia, sacral centrum, eighteen
caudal vertebrae, scapula (500 mm), coracoid, humeri (270, 265 mm), ulnae, radii,
two distal carpals, metacarpal I (50 mm), metacarpal II (90 mm), metacarpal
III, manual phalanges, manual ungual I, manual II, manual ungual III, ilium,
pubes (~460 mm), ischium (420 mm), femur (585 mm), tibiae (580 mm), fibula (560
mm), astragali, calcanea, metatarsal I (56 mm), pedal ungual I, metatarsal II,
metatarsal III (200 mm), metatarsal IV, pedal phalanges, pedal unguals (Dong,
Zhou and Zhang, 1983)
Diagnosis- (after Carrano et al., 2012) differs from Sinraptor
in- higher ratio of skull height to length (0.5); dorsal vertebral neural spines
lower (about 1.8 times centrum height); dorsal centra relatively longer; more
pronounced margin of the antorbital fossa on the jugal (also in hepingensis).
Comments- Yangchuanosaurus' holotype was discovered in 1976, while
Y. magnus'
holotype was found in September 1973. Note that early sources all give the
original description's title's translation as "Note on a new carnosaur (Yanchuangosaurus
shangyuanensis gen. et sp. nov.) from the Jurassic of Yangchuan District,
Szechuan Province." Paul (1988), Holtz et al. (2004)
and Carrano et al. (2012) all consider magnus to be synonymous with shangyouensis.
Paul believed the ilium and tibia were poorly ossified in shangyouensis'
type, showing it to be a juvenile (though far younger juveniles like Scipionyx
have expanded preacetabular processes showing full ossification). Carrano et
al. noted the (pro-?, or accessory?) maxillary fossa being developed into a
fenestra in magnus is congruent with that specimen being older. In addition,
variation in pneumatic features is common in theropods, even between sides of
the same individual. Of Dong et al.'s (1983) other characters to distinguish
the species, larger size in magnus could be ontogenetic, magnus
lacks a foramen at the base of the posterodorsal dentary process but surangular
foramina are known to vary in presence in Dromiceiomimus brevitertius
individuals, and shangyouensis' type has a tapered preacetabular process
that may be artificial but also varies in Archaeopteryx lithographica
individuals.
"Szechuanoraptor"- Dong et al. (1978) list Szechuanosaurus
"yandonensis" as a new species in a faunal list of taxa from the Wujiaba
quarry of the Shangshaximiao Formation. There is no description or illustration,
making this a nomen nudum. In 1983, Dong et al. note there was only a single
large theropod skeleton in the Wujiaba quarry, described by them as a neotype
of Szechuanosaurus campi. It can be implied that Dong et al. originally
believed CV 00214 to be a new species of Szechuanosaurus, but later decided
to include it in S. campi. Note that Dong et al.'s attempt at a neotype
designation is invalid, as the ICZN requires the original type(s) to be lost
or destroyed (Article 75.3.4) and that the new specimen "came as nearly
as practicable from the original type locality" (Article 75.3.6) whereas
CV 00214 is from a different stratigraphic group than Szechuanosaurus'
types. To make CV 00214 a neotype merely due to the suggested undiagnosability
of S. campi's syntypes would require an ICZN petition (Article 75.5).
Paul (1988) stated the teeth associated with CV 00214 lack roots and thus were
shed by a scavenger, but Dong et al. state "the tooth root is longer than
the crown, is exposed, and elliptical" for some lateral teeth. As the article
was written in Chinese and the current English tranlation dates from 1999, it
is possible Paul mistook figure 39 (which reproduces drawings of rootless S.
campi and Chienkosaurus syntypes) as figures of teeth found with CV 00214. He placed the specimen
in Metriacanthosaurus (along with Yangchuanosaurus), as Metriacanthosaurus?
sp.. Paul stated "there are so many uncertainties about this beast
that I balk at giving it a new name." Yet in the earlier discussion of
Metriacanthosaurus, Paul refers to the species as M. carpenteri
and states it belongs in a separate subgenus than M. parkeri and M.
shangyouensis. One may conclude Paul originally intended to name the taxon,
but later changed his mind and didn't catch all the times he had used the name.
In any case, "carpenteri" is a nomen nudum since ICZN Article 11.5
states "To be available, a name must be used as valid for a taxon when
proposed ..."
Chure (2000) used this specimen as the holotype of his new species "Szechuanoraptor
dongi" in his unpublished thesis. Names in theses aren't usually listed
in this website, and this one is only because it was later published by Glut
(2003). Glut's work includes a caveat to the effect that it is not available
to establish new taxonomy however, so the name remains unofficial. Chure referred
the specimens of Szechuanosaurus? zigongensis to "Szechuanoraptor
dongi" as well, but this seems incorrect, since they are from an earlier
formation and differ morphologically.
Carrano et al. (2012) agreed with Paul that CV 00214 could not be referred to
Szechuanosaurus, and also with me in that zigongensis is from
an earlier formation and they could not identify apomorphies shared between
them. They found the specimen to be sister to Yangchuanosaurus shangyouensis
in their analysis and synonymized the species.
References-
Dong, Zhang, Li and Zhou, 1978. [A new carnosaur discovered in
Yongchuan, Sichuan]. Chinese Science Bulletin. 23(5), 302-304.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (UT-CO) and a revision of the theropod family
Allosauridae. PhD thesis. Columbia University. 964 pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Glut, 2003. Dinosaurs: The Encyclopedia. Supplement 3. McFarland Press. 726
pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Y. zigongensis (Gao,
1993) Carrano, Benson and Sampson, 2012
= Szechuanosaurus zigongensis Gao, 1993
Bajocian, Middle Jurassic
Dashanpu, Xiashaximiao Formation, Sichuan, China
Holotype- (ZDM 9011) ten cervical vertebrae, cervical ribs, thirteen
dorsal vertebrae (series 1.17 m), dorsal ribs, five sacral vertebrae, twenty-five
caudal vertebrae, scapula, humerus (360 mm), radius (200 mm), ulna (240 mm),
distal carpal (46 mm), metacarpal I (62 mm), phalanx I-1 (92 mm), metacarpal
II (118 mm), phalanx II-1 (75 mm), metacarpal III (107 mm), metacarpal IV (52
mm), ilia (550 mm), pubes (580 mm), ischia (510 mm)
Referred- ?(ZDM 9012) maxilla, teeth (Gao, 1993)
?(ZDM 9013) ten teeth (Gao, 1993)
?(ZDM 9014) femur, tibia, fibula (Gao, 1993)
Diagnosis- (after Rauhut, 2000) differs from Gasosaurus and Xuanhanosaurus
in the more rectangular deltopectoral crest and the proximal part of the humerus
being less expanded transversely; from Monolophosaurus and Eustreptospondylus
in the gradually sloping anterior rim of the maxilla and the lack of opisthocoelous
cervical vertebrae; from Piatnitzkysaurus in the gradually sloping anterior
rim of the maxilla and the less expanded proximal humerus; from Poekilopleuron
in the more strongly pronounced olecranon process of the ulna and the lack of
a medial process on the radius; from Metriacanthosaurus in the less steeply
sloping posterodorsal rim of the ilium; from Proceratosaurus in the more
massive and relatively shorter posterior part of the maxilla.
(after Carrano et al., 2012) only cervical vertebrae 2-4 opisthocoelous,
remainder amphiplatyan.
Other diagnoses- Gao's (1993)
original diagnosis was- "moderate in size with an approximate length of
six meters. Cervical vertebra count is 10, with opisthocoelous anterior
and median centra and platycoelous posterior centra. Cervical ribs are
gracile and long. Dorsal vertebral count is 13, centra are
amphiplatyan, and plate-shaped neural spines are moderately high. The
five sacral neural spines are unfused. Anterior caudal neural spines
are narrow, and the prezygapophyses on the posterior caudals are
elongated. The scapula is moderately broad. The deltopectoral crest on
the humerus is particularly well developed. Radius is 56% the length of
the humerus. Metatarsal IV is vestigial. The ilium is particularly low,
the distal pubis is booted, and a muscular attachment crest on the
distal ischium is well developed."
Comments- ZDM 9011 was discovered in 1984 and incorrectly stated in Downs' English translation as being "initially described as Gasosaurus", but Google Translate provides the more logical translation of it being "a carnosaur specimen [that] was significantly different from Gasosaurus constructus". It was described by Gao (1993) as a species of Szechuanosaurus
based on a number of characters shared with CV 00214 and He's Hexi
Commune tetanurine material that are largely symplesiomorphies-
"opisthocoelous anterior and median cervicals with plate shaped neural
spines; 13 amphiplatyan dorsals with moderately high plate-shaped
neural spines that are constricted at their midsection; 5 fused
sacrals; platycoelous caudals; ilium being low, pubic shaft being
slender and long with an undeveloped distal boot; and there is an
expanded distal ischiac crest for muscle attachment". Most of
these are repeated in the diagnosis of this new species, S. zigongensis.
Chure (2000) referred the hypodigm to his new taxon "Szechuanoraptor
dongi" (based on CV 00214), needlessly creating a new species name, but
there appears to be little reason for such a referral, as Carrano et al. (2012)
note. The specimens derive from different formations and differ morphologically,
with Carrano et al. finding zigongensis to be sister to Yangchuanosaurus
shangyouensis instead. They therefore renamed it Yangchuanosaurus zigongensis.
While recovered outside Orionides by both Rauhut (2000) and Holtz et al. (2004),
this requires 11 more steps in Carrano et al.'s analysis, so is probably not
true.
References- Gao, 1993. A new species of Szechuanosaurus from the
Middle Jurassic of Dashanpu, Zigong, Sichuan. Vertebrata PalAsiatica. 31(4),
308-314.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (UT-CO) and a revision of the theropod family
Allosauridae. PhD thesis. Columbia University. 964 pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Y? sp. (Stiegler, 2019)
Late Jurassic
Nan'an, Chongqing, China
Comments- This was listed in Figure 5.2 of Stiegler's (2019) thesis as "Undescribed cf. Yangchuanosaurus, Nan’an, Chongqing, China."
Reference-
Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
Washington University. 693 pp.
Metriacanthosaurinae Paul, 1988
Definition- (Metriacanthosaurus parkeri <- Yangchuanosaurus
shangyouensis) (Hendrickx, Hartman and Mateus, 2015; modified from Carrano,
Benson and Sampson, 2012)
Diagnosis- (after Carrano et al., 2012) anteroventral border of maxillary
antorbital fossa demarcated by raised ridge; pronounced ventral keel on anterior
dorsal vertebrae (also in Carcharodontosaurus); straight posterior margin
of iliac postacetabular process; angle of less than 60 degrees between long
axes of pubic shaft and boot; ventrally curved ischial shaft; bulbous fibular
crest on tibia.
Comments- Paul (1988) originally used this as a subfamily of eustreptospondylids,
including all metriacanthosaurids known at the time (Yangchuanosaurus
and Metriacanthosaurus). Carrano et al. (2012) later recovered a much
larger Metriacanthosauridae and used Paul's subfamily as a name for one subclade,
though they mistakenly think Paul named the family instead and thus credit themselves
with the subfamily.
References- Paul, 1988. Predatory Dinosaurs of the World. Simon &
Schuster, New York. 464 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
unnamed metriacanthosaurine (Buffetaut and Suteethorn, 2007)
?Oxfordian-Early Valanginian, ?Late Jurassic-Early Cretaceous
Kham Phok, Phu Kradung Formation, Thailand
Material- (SM 10; Kham Phok metriacanthosaurid) tibia (618 mm)
Comments- Buffetaut and Suteethorn (2007) described numerous similarities
to Sinraptor and Yangchuanosaurus, referring it to Sinraptoridae (= Metriacanthosauridae).
Samathi et al. (2016) noted it has a bulbous fibular crest like
metriacanthosaurines in particular.
References- Buffetaut and Suteethorn, 2007. A sinraptorid theropod (Dinosauria:
Saurischia) from the Phu Kradung Formation of northeastern Thailand. Bulletin
de la Societe Geologique de France. 178, 497-502.
Samathi,
Chanthasit and Sander, 2016. New material of a new metriacanthosaurid
(Dinosauria, Theropoda) from the Phu Noi locality (Late Jurassic-Early
Cretaceous) of Thailand. Journal of Vertebrate Paleontology. Program
and Abstracts 2016, 217.
Shidaisaurus Wu, Currie, Dong,
Pan and Wang, 2009
S. jinae Wu, Currie, Dong, Pan and Wang, 2009
Aalenian-Bajocian, Middle Jurassic
Upper Lufeng Formation, Yunnan, China
Holotype- (LDM-LCA 9701-IV) braincase, three teeth, atlantal intercentrum,
axis, cervical vertebrae, third dorsal vertebra, fourth dorsal vertebra, fifth
dorsal vertebra, sixth dorsal vertebra, seventh dorsal vertebra, eighth dorsal
vertebra, ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra,
twelfth dorsal vertebra, thirteenth dorsal vertebra, two gastralia, sacral vertebrae
1+2, third sacral vertebra, fourth sacral vertebra, fifth sacral vertebra, first
caudal vertebra, second caudal vertebra, third caudal vertebra, ilium (620 mm),
pubes (620 mm), ischium (599 mm)
Diagnosis- (after Wu et al., 2009) supraoccipital excluded from foramen
magnum; paroccipital processes downturned at 110 degrees; large, pointed axial
epipophyses; thin and broad lamina between axial neural spine and epipophyses;
iliopubic ratio ~1.00; obturator notch absent on ischium; ischium >96% of
pubic length.
Comments- Wu et al. merely considered Shidaisaurus a tetanurine,
perhaps non-avetheropod due to the absent axial pleurocoel. Carrano et al. (2012)
included it in their tetanurine analysis and found it to be a basal metriacanthosaurine.
References- Wu, Currie, Dong, Pan and Wang, 2009. A new theropod dinosaur
from the Middle Jurassic of Lufeng, Yunnan, China. Acta Geologica Sinica. 83(1),
9-24.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Metriacanthosaurus Walker,
1964
M. parkeri (Huene, 1923) Walker, 1964
= Megalosaurus parkeri Huene, 1923
= Altispinax parkeri (Huene, 1923) Huene, 1932
Early-Middle Oxfordian, Late Jurassic
Upper Oxford Clay, England
Holotype- (OUM J.12144) three anterior dorsal vertebrae (110 mm), posterior
dorsal neural arch, partial posterior dorsal centrum, incomplete first sacral
vertebra, second sacral centrum, first caudal vertebra, nine proximal caudal
vertebrae, incomplete ilium (~765 mm), incomplete pubes, incomplete ischia,
femora (849.0 mm), proximal tibia
Diagnosis- (after Rauhut, 2000) dorsal margin of the ilium with a pronounced
kink over the posterior part of the acetabulum (may be due to preservation-
Carrano et al., 2012).
(after Carrano et al., 2012) ventral surfaces of posterior dorsal centra flat
and with breadth approximately 2/3 posterior height of centrum.
Comments- Pickering (unpublished ms) has referred the distal fibula NHMUK
40517 to Metriacanthosaurus parkeri, which was first mentioned by Lydekker
(1888) as Omosaurus? sp., then referred to Lexovisaurus durobrivensis
by Galton (1985). It was most recently called Dinosauria indet. by Maidment
et al. (2008). Benson (2009) redescribed Metriacanthosaurus in his thesis.
References- Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia
in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing
the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia,
and Proterosauria. British Museum of Natural History, London. 309 pp.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of
the Geological Society of America. 34, 449-458.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and
the origin of carnosaurs. Philosophical Transactions of the Royal Society of
London B. 248, 53-134.
Galton, 1985. British plated dinosaurs (Ornithischia, Stegosauridae). Journal
of Vertebrate Paleontology. 5, 211-254.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle d�couverte
et r�vision syst�matique: Implications phylog�n�tiques
et pal�obiog�ographiques. PhD thesis. 329 pp.
Maidment, Norman, Barrett and Upchurch, 2008. Systematics and phylogeny of Stegosauria
(Dinosauria: Ornithischia). Journal of Systematic Palaeontology. 6, 367-407.
Benson, 2009a. The taxonomy, systematics and evolution of the British
theropod dinosaur Megalosaurus. PhD thesis, University of Cambridge. [pp]
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Sinraptor Currie and Zhao, 1994
S. dongi Currie and Zhao, 1994
= Yangchuanosaurus dongi (Currie and Zhao, 1994) Gao, 1999
Late Oxfordian, Late Jurassic
Jiangjunmiao, Upper Shishugou Formation, Xinjiang, China
Holotype-
(IVPP V10600) (7.62 m) skull (900 mm), mandible (~893 mm), eleven dentary teeth
(32.29x15.33x10.2, 38x17.08x11.36, ~38.54x18.58x11.06,
39.97x19.89x10.54, 40.55x19x9.6, 38.42x19.58x9.48 mm; five lost), two
ceratobranchials (500 mm), atlas, axis (78 mm), third cervical vertebra
(74 mm), fourth cervical vertebra (85 mm), fifth cervical vertebra (80
mm), partial sixth cervical vertebra, partial seventh cervical
vertebra, partial eighth cervical vertebra, ninth cervical vertebra
(100 mm), tenth cervical vertebra (87 mm), incomplete cervical ribs
2-10, (dorsal series 1426.3 mm) first dorsal vertebra (83.5 mm), second
dorsal vertebra (94 mm), third dorsal vertebra (95.5 mm), fourth dorsal
vertebra (101.5 mm), fifth dorsal vertebra (110 mm), sixth dorsal
vertebra (115 mm), seventh dorsal vertebra (113 mm), eighth dorsal
vertebra (118 mm), ninth dorsal vertebra (113 mm), tenth dorsal
vertebra (119 mm), eleventh dorsal vertebra (120 mm), twelfth dorsal
vertebra (122 mm), thirteenth dorsal vertebra (122 mm), dorsal ribs
1-11, gastralia, first sacral vertebra (108 mm), second sacral centrum
(78.5 mm), third sacral centrum (99 mm), fourth sacral centrum (104.5
mm), partial fifth sacral vertebra, fifth sacral rib, first caudal
vertebra (77 mm), proximal caudal vertebra (103 mm), proximal caudal
vertebra (102.5 mm), proximal caudal vertebra (87.5 mm), proximal
caudal vertebra (85 mm), proximal caudal vertebra (95.5 mm), proximal
caudal vertebra (110 mm), scapulae (755 mm), sternum, proximal phalanx
I-1, metacarpal II (135 mm), phalanx II-2 (87 mm), metacarpal III (122
mm), phalanx III-1 (38 mm), manual ungual III (81 mm), metacarpal IV
(57 mm), ilia (682 mm), pubes (700 mm), ischia (650 mm), femora (876
mm), tibiae (776, 769 mm), fibulae (729, 697 mm), astragali, calcanea,
distal tarsal III, distal tarsal IV, metatarsal I (90 mm), phalanx I-1
(66 mm), pedal ungual I (66 mm), metatarsal II (360 mm), phalanx II-1
(135 mm), phalanx II-2 (107 mm), pedal ungual II (111 mm), metatarsal
III (410 mm), phalanx III-1 (135 mm), phalanx III-2 (98 mm), phalanx
III-3 (74 mm), pedal ungual III (90 mm), metatarsal IV (375 mm),
phalanx IV-1 (98 mm), phalanx IV-2 (82 mm), phalanx IV-3, pedal ungual
IV (86 mm), metatarsal V (65 mm)
Paratype- (IVPP coll.) nine teeth
Late Oxfordian, Late Jurassic
Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Referred- (IVPP V15310) (~12 m) anterior maxillary tooth (92x35x18 mm) (Xu and Clark, 2008)
?(IVPP V18060) (~7.2 m) metatarsal IV (355 mm) (He, Clark and Xu, 2013)
(IVPP coll.) skull, cervical vertebrae (Clark et al., 2002)
Diagnosis- (after Currie and Zhao, 1994) differs from S? hepingensis
in- longer, lower premaxilla; more numerous and elaborate accessory maxillary
fossae; posterior postorbital process with reduced lateral exposure; longer
subtemporal bar.
Comments- Grady (1993) states the holotype was found on September 2 1987
at Quarry 6, when Braman "found a small theropod near the spot where he
had been excavating a crocodile" ... and Currie "dug out two vertebrae,
a scapulocoracoid or shoulder blade, a toe bone, and the end of what
looked like a fibula." "Before the day ended three ribs, six
phalanges, a humerus, four metatarsals, and a claw nearly eight
centimeters long had been added to the list of specimens", "another,
smaller [than Sinraptor's
paratypes] allosaurid tooth" and that Currie "also found a calcaneum
and astraglus ... that ... fit together perfectly." On September 14
Currie realized "What he thought was a fibula was in fact part of a
femur" and "he had exposed two more metatarsals, two phalanges, a
distal tarsal, and the end of a fibula." It was first announced
in Anderson (1987a) as "Jiangjunmiaosaurus", a name generally associated
with what was eventually named Monolophosaurus. The article mentions
preserved feet and "the whole" skeleton, unlike the Monolophosaurus
type. Anderson's (1987b) later article clarifies that "it has been tentatively
labelled a jiangjunmiaosaurus, a carnivore first found in the Junggar Basin
in 1983 by Zhao Xi Lin." The latter matches the discovery of Monolophosaurus'
holotype, so it's apparent Sinraptor was originally thought to be another
Monolophosaurus specimen when first discovered. The second article contains
photos of Sinraptor dongi's hindlimb elements and mentions Currie felt
it was probably related to allosaurs. Paul (1988) mentions the specimen as an
allosaurine where "the nasal ridges and orbital horn are combined into
a prominent pair of long, low, rugose-surfaced crests", which clearly described
Sinraptor though Paul uses the same "Jinagjunmiaosaurus" and
mentions pers. comm. with Dong, so the specimens were probably still thought
to be conspecific. Currie and Zhao (1991) reported on both Monolophosaurus
and Sinraptor in their abstract, still unnamed, so that Olshevsky (1991)
listed both "Jiangjunmiaosaurus" and "Monolophosaurus" as
separate genera, not realizing the proper connection noted above. Dong (1992) lists this specimen as "Yangchuanosaurus new species Currie and Zhao (in press.)", and calls it "a nearly complete skeleton of a theropod that is similar to Allosaurus." The taxon was finally described
by Currie and Zhao (1994) in a monograph which assigned it to their new family
Sinraptoridae. Note that while volume 30(10) of the Canadian Journal of Earth
Sciences lists its date as October 1993, it was not published until February
or March of 1994. The quadrate was later described by Currie (2006), while the
braincase was redescribed in depth by Paulina Carabajal and Currie (2012) and the teeth by Hendrickx et al. (2020).
Currie and Zhao (1994) mention "nine isolated, shed teeth in the
collections of the IVPP that were recovered from a sauropod (IVPP
87001) found in a quarry less than a kilometre from the holotype"
"collected by the Dinosaur Project in 1987 and 1988." Notably,
Figure 65 of Dong (1992) shows that IVPP V87001 is a locality at
Jiangjunmiao, not a specimen number of the sauropod itself. The
specimen is not Mamenchisaurus? sinocanadorum (IVPP locality 87004) or Bellusaurus (IVPP locality 830003), nor is it Klamelisaurus
which was excavated by 1984. Grady (1993) writes about this
"second sauropod quarry" and notes on September 1 1987 Russell, Tang
and Danis "dislodged four carnivore teeth" that were "similar to Allosaurus
teeth: curved and barbed and containing "blood grooves" down the sides"
and that on September 2 "More teeth were found in the second sauropod
quarry: a total of nine so far, all apparently allosaurid."
Clark et al.'s (2002) SVP presentation announced a new skull and cervicals of Sinraptor, but this has not yet been published.
Xu and Clark (2008) described a large tooth found between 2001 and 2007
"preserved within a partial sauropod skeleton." as "a gigantic
sinraptorid, either a taxon assignable or closely related to Yangchuanosaurus magnus [seemingly based on size alone], ... or an old individual of Sinraptor." Its crown length is 145% the size of S. dongi's
largest preserved holotype tooth (rmax4), and its Crown Base Ratio is
almost identical to the third maxillary tooth and an isolated dentary
tooth (0.51 vs. 0.5043 and 0.5053) which it is 155% and 227% the size
of respectively. The Crown Height Ratio of 2.63 is more similar to the
third maxillary (2.4371) than the dentary tooth (2.1342) though,
suggesting it is an anterior maxillary tooth as well. Using the
155% ratio would lead to a body length of 11.8 meters, rounded to 12
here. Denticle morphology appears identical. Size
differences between it and the type could easily be ontogenetic or
individual.
He et al. (2013)
described a metatarsal IV from Wucaiwan discovered since 2000 by the
joint IVPP-CAS-GWU expedition, resolving it sister to Sinraptor dongi in their phylogenetic analysis of metatarsal IV characters. They
noted it resembled S. dongi except for being more robust and having a
lateral condyle smaller than the medial one. While the authors interpreted these
as species-level differences, they are here considered individual variation
considering the vast amount of variation in e.g. Allosaurus and Yangchuanosaurus. It should be noted that besides the S. dongi holotype the only preserved sinraptorid metatarsal IV is that of Yangchuanosaurus
specimen CV 00214, which was described in very basic terms and only
figured in oblique medial view largely covered by other pedal elements.
References- Anderson, 1987a. Chinese dinosaur dig strikes bonanza. New
Scientist. 1584, 25.
Anderson, 1987b. Chinese unearth a dinosaurs' graveyard. New Scientist. 1586,
28-29.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Currie and Zhao, 1991. Two new theropods from the Jurassic of Xinjiang, People's
Republic of China. Journal of Vertebrate Paleontology. 11(3), 24A.
Dong, 1992. Dinosaurian Faunas of China. Ocean Press/Springer-Verlag.
188 pp.
Grady, 1993. The Dinosaur Project: The Story of the Greatest
Dinosaur Expedition Ever Mounted. Macfarlane
Walter & Ross and The Ex Terra Foundation. 261 pp.
Currie and Zhao, 1994. A new carnosaur (Dinosauria, Theropoda) from the Jurassic
of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences.
30(10), 2037-2081.
Gao, 1999. A complete carnosaur skeleton from Zigong, Sichuan. Sichuan Science
& Technology Press, Chengdu. 80 pp.
Clark, Xu, Forster, Wang and Andres, 2002. New small dinosaurs from the Upper
Jurassic Shishugou Formation at Wucaiwan, Xinjiang, China. Journal of Vertebrate
Paleontology. 22(3), 44A.
Currie, 2006. On the quadrate of Sinraptor dongi (Theropoda: Allosauroidea)
from the Late Jurassic of China. In Csiki (ed.). Mesozoic and Cenozoic Vertebrates
and Paleoenvironments. Ars Docendi. 111-115.
Xu and Clark, 2008. The presence of a gigantic theropod in the Jurassic Shishugou
Formation, Junggar Basin, Western China. Vertebrata PalAsiatica. 46(2), 157-160.
Paulina Carabajal and Currie, 2012. New information on the braincase of Sinraptor
dongi (Theropoda: Allosauroidea): Ethmoidal region, endocranial anatomy,
and pneumaticity. Vertebrata PalAsiatica. 50(2), 85-101.
He, Clark and Xu, 2013. A large theropod metatarsal from the upper part of Jurassic
Shishugou Formation in Junggar Basin, Xinjiang, China. Vertebrata PalAsiatica.
51(1), 29-42.
Hendrickx, Stiegler, Currie, Han, Xu, Choiniere and Wu, 2020. Dental anatomy of the apex predator Sinraptor dongi (Theropoda: Allosauroidea) from the Late Jurassic of China. Canadian Journal of Earth Sciences. 57(9), 1127-1147.
S? hepingensis (Gao, 1992)
Currie and Zhao, 1994
= Yangchuanosaurus hepingensis Gao, 1992
Bathonian-Callovian, Middle Jurassic
Tianwan, Shangshaximiao Formation, Sichuan, China
Holotype- (ZDM 0024) (8.84 m) skull (1.04 m), stapes, mandibles (1 m),
proatlas, atlantal intercentrum (36 mm), atlantal neural arch, axis (88 mm),
third cervical vertebra (88 mm), fourth cervical vertebra (98 mm), fifth cervical
vertebra (100 mm), sixth cervical vertebra (113 mm), seventh cervical vertebra
(110 mm), eighth cervical vertebra (115 mm), ninth cervical vertebra (115 mm),
incomplete tenth cervical vertebra (100 mm), cervical ribs, incomplete first
dorsal vertebra (95 mm), second dorsal vertebra (85 mm), third dorsal vertebra
(90 mm), fourth dorsal vertebra (99 mm), fifth dorsal vertebra, sixth dorsal
vertebra, seventh dorsal vertebra (115 mm), eighth dorsal vertebra (115 mm),
ninth dorsal vertebra (115 mm), tenth dorsal vertebra (121 mm), eleventh dorsal
vertebra (123 mm), twelfth dorsal vertebra (127 mm), thirteenth dorsal vertebra
(123 mm), complete to incomplete dorsal ribs (50-960 mm), gastralium, sacrum
(120, 118, 105, 105, 128 mm), first caudal vertebra, second caudal vertebra
(106 mm), third caudal vertebra (106 mm), fourth caudal vertebra (105 mm), fifth
caudal vertebra (108 mm), sixth caudal vertebra (110 mm), seventh caudal vertebra
(118 mm), eighth caudal vertebra (123 mm), ninth caudal vertebra (125 mm), tenth
caudal vertebra (126 mm), eleventh caudal vertebra (126 mm), twelfth caudal
vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra (115 mm), fifteenth
caudal vertebra (114 mm), sixteenth caudal vertebra (115 mm), seventeenth caudal
vertebra (112 mm), eighteenth caudal vertebra (110 mm), nineteenth caudal vertebra
(105 mm), twentieth caudal vertebra (104 mm), twenty-first caudal vertebra (103
mm), twenty-second caudal vertebra (103 mm), twenty-third caudal vertebra (100
mm), twenty-fourth caudal vertebra (100 mm), twenty-fifth caudal vertebra, twenty-sixth
caudal vertebra (98 mm), twenty-seventh caudal vertebra (97 mm), twenty-eighth
caudal vertebra (90 mm), twenty-ninth caudal vertebra, thirtieth caudal vertebra
(85 mm), thirty-first caudal vertebra (82 mm), thirty-second caudal vertebra
(80 mm), thirty-third caudal vertebra (75 mm), thirty-fourth caudal vertebra
(70 mm), ten chevrons, scapulae (760, 740 mm), coracoids (250 mm), ilia (760,
760 mm), pubes (795, 805 mm), ischia (770, 750 mm), femur (980 mm)
Diagnosis- (after Carrano et al., 2012) differs from Yangchuanosaurus
in- proportionally long, low skull; very tall dorsal vertebral neural spines.
differs from Sinraptor dongi in- sinuous, rugose nasal crest; marked
margin of jugal antorbital fossa; more horizontally oriented pubic boot.
Comments- The holotype was discovered in 1985 and initially thought to
be a specimen of Yangchuanosaurus shangyouensis (e.g. fig. 57 in Dong,
1992 which labels it "A new, nearly complete skeleton of Yangchuanosaurus shangyouensis was collected by Zigong Dinosaur Museum from the vicinity (Heping) of Zigong in 1986"). While initially assigned to Yangchuanosaurus in its description
(Gao, 1992), Currie and Zhao (1994) placed hepingensis in their new genus
Sinraptor. Note that while volume 30(10) of the Canadian Journal of Earth
Sciences lists its date as October 1993, it was not published until February
or March of 1994. Gao (1999) and Xing et al. (2009) disagree, placing both hepingensis
and dongi in Yangchuanosaurus. Carrano et al. (2012) found hepingensis,
Shidaisaurus, Siamotyrannus and Metriacanthosaurus to be
closer to Sinraptor than to Yangchuanosaurus. Thus Sinraptor
cannot merely be synonymized with Yangchuanosaurus without including
all metriacanthosaurids, in which case Metriacanthosaurus would have
priority.
References- Dong, 1992. Dinosaurian Faunas of China. Ocean Press/Springer-Verlag,
Beijing/Berlin. 188 pp.
Gao, 1992. Yangchuanosaurus hepingensis, a new species of carnosaur from
Zigong, Sichuan. Vertebrata PalAsiatica. 30, 313-324.
Currie and Zhao, 1994. A new carnosaur (Dinosauria, Theropoda) from the Jurassic
of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences.
30(10), 2037-2081.
Gao, 1999. A complete carnosaur skeleton from Zigong, Sichuan. Sichuan Science
& Technology Press, Chengdu. 80 pp.
Xing, Dong, Peng, Shu, Hu and Jiang, 2009. A scapular fracture in Yangchuanosaurus
hepingensis (Dinosauria: Theropoda). Geological Bulletin of China. 28(10),
1390-1395.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Allosauridae sensu Padian and Hutchinson, 1997
Definition- (Allosaurus fragilis <- Sinraptor dongi) (modified)
Allosauria Paul, 1988
Definition- (Allosaurus fragilis + Carcharodontosaurus saharicus,
-Metriacanthosaurus parkeri) (Hendrickx, Hartman and Mateus, 2015)
Other definition- (Allosaurus fragilis + Carcharodontosaurus saharicus) (Rauhut and Pol, 2019)
= Allosauria sensu Rauhut and Pol, 2019
Definition- (Allosaurus fragilis + Carcharodontosaurus saharicus)
Comments- Paul (1988) used Allosauria for what would now be called Carnosauria,
though he excluded metriacanthosaurids and included Ornitholestes, Proceratosaurus
and tyrannosaurids. As Paul's Allosauridae included known carcharodontosaurids
(Shaochilong, Acrocanthosaurus, Altispinax, Labocania
and Carcharodontosaurus), Carrano et al. (2012) used Allosauria for the
allosaurid+carcharodontosaurid clade of carnosaurs. They provided no definition,
though (Allosaurus fragilis + Carcharodontosaurus saharicus, -Metriacanthosaurus
parkeri) would be the obvious choice and was later published by Hendrickx
et al. (2015). Several analyses such as Allain (2002) have instead found carcharodontosaurids
to group with metriacanthosaurids to the exclusion of allosaurids, which takes
six more steps when enforced in Carrano et al.'s matrix, so is still quite possible.
References- Paul, 1988. Predatory Dinosaurs of the World. Simon &
Schuster, New York. 464 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle
Jurassic Ca�ad�n Asfalto Formation of Argentina highlights phylogenetic
uncertainty in tetanuran theropod dinosaurs. Scientific Reports.
9:18826.
Carcharodontosauridae Stromer, 1931
Definition- (Carcharodontosaurus saharicus <- Allosaurus fragilis,
Sinraptor dongi) (Holtz et al., 2004)
Other definitions- (Carcharodontosaurus saharicus <- Allosaurus
fragilis, Sinraptor dongi, Monolophosaurus jiangi, Cryolophosaurus
ellioti) (modified from Sereno, 1998)
(Carcharodontosaurus saharicus <- Allosaurus fragilis, Sinraptor
dongi, Passer domesticus) (Brusatte and Sereno, 2008)
(Carcharodontosaurus saharicus <- Neovenator salerii, Allosaurus
fragilis, Sinraptor dongi) (Benson, Carrano and Brusatte, 2010)
= "Acrocanthosauridae" Molnar, 2001
= Carcharodontosauridae sensu Sereno, 1998
Definition- (Carcharodontosaurus saharicus <- Allosaurus fragilis,
Sinraptor dongi, Monolophosaurus jiangi, Cryolophosaurus ellioti)
(modified)
= Carcharodontosauridae sensu Brusatte and Sereno, 2008
Definition- (Carcharodontosaurus saharicus <- Allosaurus fragilis,
Sinraptor dongi, Passer domesticus)
= "Carcharodontosauria" Benson, Carrano and Brusatte, 2009 online
= Carcharodontosauria Benson, Carrano and Brusatte, 2010
Definition- (Carcharodontosaurus saharicus, Neovenator salerii
<- Allosaurus fragilis, Sinraptor dongi) (Benson, Carrano and
Brusatte, 2010)
Comments- Brusatte and Sereno's (2008) definition differs from Holtz
et al.'s (2004) by including Passer as an external specifier. The only
times carcharodontosaurids have been placed in Coelurosauria is when tyrannosaurids
were as well (Bakker et al., 1988; Paul, 1988), and they have often been placed
closer to tyrannosaurids than to Allosaurus or Passer (Paul, 1988;
Kurzanov, 1989; Molnar et al., 1990). So Tyrannosaurus would be a more
useful tertiary external specifier than Passer. The remote possibility
of a relationship to ceratosaurs (Bonaparte et al., 1990) might suggest Carnotaurus
should be used as an additional external specifier.
Molnar (2001) proposed Acrocanthosauridae for at least Acrocanthosaurus
and Carcharodontosaurus, but did not provide a definition or diagnosis
(ICZN Article 13.1.1), so the taxon is a nomen nudum.
Benson et al. (2010) found megaraptorans grouped with Neovenator as basal
carcharodontosaurids, but redefined Carcharodontosauridae to include only those
taxa closer to Carcharodontosaurus than to Neovenator. They then
gave their new taxon Carcharodontosauria the tradition definition for Carcharodontosauridae.
These taxonomic changes are here seen as unecessary and not followed. Benson
et al.'s paper was originally released online in November 2009 but not officialy
published until January 2010.
References- Molnar, 2001. Theropod paleopathology: A literature survey.
In Tanke and Carpenter (eds). Mesozoic Vertebrate Life, new research inspired
by the paleontology of Philip J. Currie. Indiana University Press. 337-363.
Benson, Carrano and Brusatte, 2010. A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Lusovenator Malafaia, Mocho, Escaso and Ortega, 2020
L. santosi Malafaia, Mocho, Escaso and Ortega, 2020
Late Kimmeridgian, Late Jurassic
Praia de Valmit�o, Praia da Amoreira-Porto Novo Formation, Portugal
Holotype-
(SHN.036/1-23, 25-29, 33-35, 37, 40-45, 50-68, 70-72, 80-90) (~3-3.5 m; juvenile) atlantal neurapophyses, atlantal
intercentrum (15.7 mm),
odontoid (14.15 mm), partial cervical vertebra, four cervical
prezygapophyses, four
cervical postzygapophyses, three cervical neural spines, several
cervical rib fragments, two partial anterior dorsal vertebrae (58.58,
~47.44 mm), eight
partial mid to posterior dorsal centra (49.12, 41.83, 56.13, 55.26,
40.04, 43.6 mm), several mid to posterior dorsal
prezygapophyses, five mid to posterior dorsal neural spines, mid to
posterior central and neural arch fragments, two partial mid dorsal
ribs, proximal posterior dorsal rib, several rib fragments,
incomplete first sacral centrum (47.21 mm), two partial mid sacral
centra (65.4 mm),
incomplete fifth sacral centrum (57 mm), partial fifth sacral neural
arch,
sacral neural spine fragment, incomplete first caudal vertebra (61.07
mm), three proximal caudal centra (61.21, 59.16, 60.19 mm), partial
~twentieth-twenty-eighth caudal vertebrae (60.12, 63.93, 63.2, 66.39,
60.88, 60.73, 68.62, 66.37 mm), several caudal transverse process
fragments, caudal zygapophyseal fragments, caudal neural spine
fragment, seven incomplete proximal to mid chevrons, several chevron
fragments, ilia (one incomplete, one fragmentary (420 mm), pubes (330
mm),
ischia (265, 270 mm)
Late Tithonian, Late Jurassic
Praia de Cambelas, Freixial Formation, Portugal
Paratype-
(SHN.019/2-13) two proximal caudal centra (82.6, 89.8 mm), proximal caudal
centrum fragments, eighteenth caudal vertebra (90.0 mm), nineteenth
caudal vertebra (91.0 mm), twentieth caudal vertebra (95.2 mm),
twenty-first caudal vertebra (91.1 mm), twenty-second caudal vertebra
(100.4 mm), twenty-third caudal vertebra (99.1 mm), twenty-fourth
caudal vertebra (96.1 mm), twenty-fifth caudal vertebra (93.1 mm),
twenty-sixth caudal vertebra (95.4 mm), twenty-seventh caudal vertebra
(94.5 mm), twenty-eighth caudal vertebra (89.7 mm), twenty-ninth caudal
vertebra (89.3 mm), thirtieth caudal vertebra (79.2 mm), thirty-first
caudal vertebra (86.9 mm), thirty-second caudal vertebra (84.4 mm),
thirty-third caudal vertebra (80.4 mm), thirty-fourth caudal vertebra
(78.5 mm), fragmentary to complete nineteenth-thirty-second chevrons,
fragmentary femur, tibial shaft fragment, incomplete fibula, calcaneum,
partial distal tarsal III, distal tarsal IV, metatarsal I (95.53 mm),
phalanx I-1 (75.96 mm), pedal ungual I (70.01 mm), distal metatarsal
II, phalanx II-1 (103.27 mm), phalanx II-2 (88.82 mm), pedal ungual II
(113.42 mm), incomplete metatarsal III, phalanx III-1 (112.11 mm),
phalanx III-2 (89.89 mm), phalanx III-3 (71.97 mm), pedal ungual III
(90.74 mm), incomplete metatarsal IV (~326 mm), phalanx IV-1 (84.8 mm),
phalanx IV-2 (71.88 mm), phalanx IV-3 (58.33 mm), phalanx IV-4 (40.67
mm), pedal ungual IV (74.92 mm), metatarsal V (~106.06 mm) (Malafaia et
al., 2009)
Late Kimmeridgian-Early Tithonian, Late Jurassic
Porto das Barcas, Sobral Formation, Portugal
Referred- ?(ML 555) femur (Antunes and Mateus, 2003)
Diagnosis- (after Malafaia et
al., 2020) large recesses in neural arch of anterior dorsal vertebrae;
well-developed and continuous longitudinal laminae extending from the
tip of the prezygapophyses to the distal end of the postzygapophyses in
mid caudal vertebrae; supraacetabular crest of ilium forming a
prominent ventrolaterally projecting shelf.
Comments- The
holotype was found by a
private collector and donated to the SHN in 2008. As noted by
Malafaia et al. (2017), "three isolated teeth were collected in the
same site and they have a morphology and size compatible with the
postcranial elements", but they "do not preserve any parts of the root,
suggesting that they correspond to shed teeth." They thus
excluded the teeth (SHN.03630-32) from the hypodigm. Malafaia et al.
(2008) referred gastralia to the specimen, but these are not mentioned
or listed in the 2017 osteology. Malafaia et al. (2020) reidentify the
supposed articulated axis and axial intercentrum as a partial
post-axial cervical vertebra.
The holotype was originally described in an extended abstract by
Malafaia et al. (2008) as Allosauroidea indet., though they noted
differences from both Lourinhanosaurus and Allosaurus. They further said "it
remains to be determined whether the differences detected in the
Valmit�o theropod are due to the specimen's belonging to a new taxon,
or, more likely, indicate that it is an immature specimen of the
previously described taxa." Malafaia et al. (2017) described
the specimen in detail as Allosauroidea gen. et sp. indet. (despite
recovering four autapomorphies), and added it to Carrano et al.'s
tetanurine matrix to find it resolved as a non-carcharodontosaurid
allosaurian. Malafaia et al. (2009) first described SHN.019 in a 2007
symposium proceeding as "consistent with the available material of Allosaurus fragilis from Andr�s, therefore they are preliminarily considered co-specific." It
was later described in more detail by Malafaia et al. (2019) as
Carcharodontosauria gen. et sp. indet., who stated "this specimen
together with a previously described specimen, SHN.036, may represent
an allosauroid distinct from the taxa currently known in the Upper
Jurassic of the Lusitanian Basin. Malafaia et al. (2020) followed this
to its conclusion, naming a new taxon using SHN.036 as the holotype
with SHN.019 referred, finding Lusovenator to be a basal carcharodontosaurid in a trichotomy with Siats and more derived taxa, again based on a version of Carrano et al.'s analysis.
A femur (ML 555) from the Sobral Formation originally referred to Lourinhanosaurus
by Antunes and Mateus (2003) was removed from that taxon by Malafaia et
al. (2020). They stated "ML 555 has a femoral head that projects
dorsomedially, which is a feature that has been interpreted as a
synapomorphy of Carcharodontosauria", so it is here provisioonally
referred to Lusovenator although it cannot be directly compared to the holotype and has not been compared to the paratype.
References-
Antunes and Mateus, 2003. Dinosaurs of Portugal. Comptes Rendus Palevol. 2(1), 77-95.
Malafaia, Ortega, Silva, Escaso and Dantas, 2008. Un nuevo
ejemplar de Allosauroidea (Dinosauria: Tetanurae) del Jur�sico Superior
de Valmit�o (Lourinh�, Portugal). XXIV Jornadas de la Sociedad
Espa�ola de Paleontolog�a, abstracts. 148-149.
Malafaia, Ortega, Escaso, Silva, Ramalheiro, Dantas, Moniz and Barriga, 2009.
A preliminary account of a new Allosaurus individual from the Lourinh�
Group (Upper Jurassic of Torres Vedras, Portugal). Actas de las IV Jornadas
Internacionales sobre Paleontolog�a de Dinosaurios y su Entorno. 243-251.
Malafaia, Mocho, Escaso and Ortega, 2017 (online 2016). A juvenile
allosauroid theropod (Dinosauria, Saurischia) from the Upper Jurassic
of Portugal. Historical Biology. 29, 654-676.
Malafaia, Mocho, Escaso, Dantas and Ortega, 2019 (online 2018).
Carcharodontosaurian remains (Dinosauria, Theropoda) from the Upper
Jurassic of Portugal. Journal of Paleontology. 93, 157-172.
Malafaia, Mocho, Escaso and Ortega, 2020. A new carcharodontosaurian
theropod from the Lusitanian Basin: Evidence of allosauroid sympatry in
the European Late Jurassic. Journal of Vertebrate Paleontology.
e1768106.
undescribed carcharodontosaurid (Calvo et al., 2004)
Albian, Early Cretaceous
Gorro Frigio Formation, Chubut, Argentina
Material- (MEF 1157) cervicals, caudals, scapula
Reference- Calvo, Porfiri, Veralli, Novas and Pobletei, 2004. Phylogenetic
status of Megaraptor namunhuaiquii Novas based on a new specimen from
Neuqu�n, Patagonia, Argentina. Ameghiniana. 41(4), 565-575.
unnamed carcharodontosaurid (Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016)
Middle Cenomanian-Early Turonian, Late Cretaceous
Violante farm, Huincul Formation of the Rio Limay Subgroup, Rio Negro, Argentina
Material-
(MPCA-Pv 803) posterior dorsal centrum, fragmentary rib, distal tarsal
III, distal tarsal IV, metatarsal II (435 mm), metatarsal III, pedal ungual ?III, distal
?metatarsal fragment, phalanx IV-1, phalanx IV-3 (60 mm), pedal
ungual ?IV
Comments- Motta et al. identify this as Carcharodontosauridae gen. et sp. indet.. Note it may belong to Mapusaurus
from the same locality. The curved and less transversely expanded
proximal end of metatarsal II with a beveled proximal surface resemble Mapusurus more than Meraxes. Based on the near complete pes of Meraxes, the pedal ungual in Figure 14 is identified here as III and that in Figure 15 as IV.
Reference- Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016. New theropod fauna from the Upper Cretaceous (Huincul
Formtation) of northwestern Patagonia, Argentina. In Khosla and Lucas (eds.).
Cretaceous period: Biotic diversity and biogeography. New Mexico Museum of Natural
History and Science Bulletin. 71, 231-253.
Neovenatoridae Benson, Carrano and Brusatte, 2010
Definition- (Neovenator salerii <- Carcharodontosaurus saharicus,
Allosaurus fragilis, Sinraptor dongi) (Benson, Carrano and Brusatte, 2010)
= "Neovenatoridae" Benson, Carrano and Brusatte, 2009 online
Comments- Benson et al.'s paper was originally released online in November
2009 but not officially published until January 2010.
Reference- Benson, Carrano and Brusatte, 2010. A new clade of archaic
large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to
the latest Mesozoic. Naturwissenschaften. 97(1), 71-78.
Neovenator Hutt, Martill and Barker,
1996
= "Neovenator" Naish, 1996
N. salerii Hutt, Martill and Barker, 1996
= "Neovenator salerii" Naish, 1996
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (MIWG 6348) (~7.5 m; subadult or adult) (skull ~798 mm) premaxillae (87 mm), incomplete maxilla, incomplete
nasal, incomplete palatine, anterior dentary (lost), teeth, axis (58 mm), partial
sixth cervical vertebra (76 mm), eighth cervical vertebra (66 mm), incomplete
ninth cervical vertebra (75 mm), proximal cervical rib, ninth dorsal vertebra
(92 mm), incomplete tenth dorsal vertebra (95 mm), fourteenth dorsal vertebra
(110 mm), two partial dorsal ribs, rib fragments, ten-fifteen fragmentary gastralia,
partial first caudal vertebra (109 mm), second caudal vertebra (99 mm), incomplete
third caudal vertebra (105 mm), incomplete fourth caudal vertebra (109 mm),
incomplete fifth caudal vertebra (107 mm), incomplete sixth caudal vertebra
(107 mm), incomplete seventh(?) caudal vertebra (106 mm), ninth(?) caudal neural
arch, incomplete fourteenth(?) caudal vertebra (103 mm), incomplete seventeenth(?)
caudal vertebra (103 mm), eighteenth(?) caudal centrum (102 mm), incomplete
twenty-first(?) caudal vertebra (94 mm), incomplete twenty-second(?) caudal
vertebra (95 mm), fused twenty-fifth(?) (91 mm) and twenty-sixth caudal vertebra
(92 mm) and twenty-sixth chevron, incomplete twenty-seventh(?) caudal vertebra
(88 mm), twenty-eighth(?) caudal vertebra (81 mm), incomplete distal caudal
vertebra (79 mm), incomplete distal caudal vertebra (77 mm), distal caudal vertebra
(75 mm), distal caudal vertebra (73 mm), distal caudal vertebra (71 mm), incomplete
distal caudal vertebra (68 mm), partial proximal chevron, two mid chevrons (~100
mm), distal chevron, incomplete scapula (~505 mm), incomplete coracoid (140
mm), proximal ischium, femora (730, 730 mm), tibiae (680, 685 mm), fibula (618
mm), metatarsal II (315 mm), phalanx II-1 (109 mm), phalanx II-2 (88 mm), proximal
metatarsal III (~343 mm), pedal unguals III (one partial; 117 mm), metatarsal
IV (325 mm), phalanx IV-1 (83 mm), phalanx IV-2 (69 mm), phalanges IV-3 (52
mm), phalanx IV-4 (34 mm), pedal ungual IV (82 mm), metatarsal V (150 mm)
....(NHMUK R10001) teeth, incomplete
fifth cervical vertebra (62 mm), seventh cervical vertebra (73 mm), first dorsal
vertebra (71 mm), incomplete second dorsal vertebra (62 mm), fourth dorsal transverse
processes, partial fifth dorsal vertebra (87 mm), incomplete sixth dorsal vertebra
(77 mm), incomplete seventh dorsal vertebra (93 mm), incomplete eighth dorsal
vertebra (89 mm), incomplete twelfth dorsal vertebra (115 mm), partial thirteenth
dorsal vertebra (117 mm), three or four sacral centra, caudal vertebrae, fragmentary
ilia, proximal pubis, pubic shaft fragments, ischial shaft fragments, distal
ischia
Paratype- (MIWG 6352) (subadult) second dorsal centrum, fifth dorsal
centrum (~77 mm), partial seventh dorsal neural arch, several rib fragments,
fused second-fourth sacral centra, first or fifth sacral centrum, sacral neural
arch, partial chevron, axial fragments, partial ilium, pubes (560 mm), proximal
ischium
Referred- ?(Dinosaur Expeditions Centre coll.) incomplete proximal caudal
vertebra (Mattsson pers. comm., 2015)
?(Dinosaur Expeditions Centre coll.) distal tibia (Mattsson pers. comm., 2015)
?(IWCMS 2000.1108) distal ____ (Brusatte et al., 2008)
?(IWCMS 2002.186) vertebral fragments including two anterior cervical neural
arches, long bones (Brusatte et al., 2008)
?(MIWG 4199) (~10 m) pedal phalanx (Hutt, 2001)
?(MIWG 5121) tooth (Brusatte et al., 2008)
(MIWG 5470) (subadult or adult) incomplete ninth cervical neural arch, incomplete
eighth dorsal vertebra, pedal phalanx III-1 (Hutt, 2001)
(MIWG 6348) proximal caudal vertebra (Hutt, 2001)
Diagnosis- (after Hutt et al., 1996) five premaxillary teeth (also in
Allosaurus); external naris twice as long as high; maxilla with large
maxillary fenestra approximately one sixth the length of the maxillary tooth
row; tooth crowns one quarter total tooth length; pedal unguals with groove
on extensor surface.
(after Naish et al., 2001) premaxilla with accessory interpremaxillary peg and
socket articulation in the dorsal region of the symphysis.
(after Brusatte et al., 2008) transverse expansion of the anterior articular
surface of the axial intercentrum; lateral foramina on the anterior surface
of the odontoid; small single foramen on the lateral surface of the axial neural
spine; fusion of the cervical ribs to the posterior cervical vertebrae; camellate
internal texture exposed externally on parapophyseal facets of eighth and ninth
cervical vertebrae (possibly due to fusion with ribs); ventral surfaces of anterior
dorsal vertebrae developed as sharp ridges, not inset from the lateral surface
of the centra; hypapophyses of anterior dorsal vertebrae developed as low mound-like
eminences; curved flanges emerging laterally from pre- and postzygapophyseal
facets of posterior dorsal vertebrae; scapula-coracoid glenoid fossa that is
wider mediolaterally than long anteroposteriorly; shelf adjacent to the preacetabular
notch on the medial surface of the ilium; distal ischial boot in which the left
and right ischia are conjoined anteriorly but diverge posterolatcrally; femoral
head oriented anteromedially and inclined proximally; robust ridge on the external
surface of the lesser trochanter of the femur; thumbprint-shaped depression
on posterior surface of femoral shaft lateral to the proximal end of the fourth
trochanter; proximodistally short, notch-like extensor groove and almost flat
anterior surface of distal end of femur; suboval rugosity on medial surface
of distal tibia; anteroposteriorly pinched proximal portion of the lateral malleolus
of the distal tibia; ventral spine on anterolateral crest of fibular condyle
of tibia; concave lateral surface of metatarsal II for articulation with metatarsal
III.
Other diagnoses- Hutt et al. (1996) included a large external naris in
their diagnosis, but Brusatte et al. (2008) noted it was similar in size to
most carnosaurs. Posterior dorsal centra that are pleurocoelous are found in
other carcharodontosaurids as well.
Naish et al. (2001) included high dorsolateral nasal crests in their diagnosis,
but Brusatte et al. (2008) note these are also present in Allosaurus.
Brusatte et al. also note serrations which complete across tooth tips are now
known in many other theropod taxa.
Comments- The holotype was discovered in 1978 and collected over the
next two decades, becoming accessioned in two museums. Hutt (1999) and Martill
and Naish (2001) referred to the dentary in the holotype as BMNH R10001, but
it has apparently never been stored at the NHMUK (was BMNH) and may be an MIWG specimen.
The paratype was discovered in 1987 and originally thought to be a different
species (Hutt et al., 1990), as the holotype was thought to have a small pubic
boot based on an incorrectly identified ischium. MIWG 5470 was discovered in
1985.
Though generally believed to be a basal carcharodontosaurid now, Neovenator
was originally assigned to Allosauridae. This requires 7 more steps when enforced
in Carrano et al.'s (2012) matrix, so is less likely but still possible.
References- Hutt, Simmonds and Hullman, 1990. Predatory dinosaurs from
the Isle of Wight: Proceedings of the Isle of Wight Natural History and Archaeological
Society. 9, 137-146.
Naish, 1996. Isle of Wight. Dinosaur Discoveries. 1, 15.
Hutt, Martill and Barker, 1996. The first European allosauroid dinosaur (Lower
Cretaceous, Wealden Group, England). Neues Jahrbuch f�r Geologie und Pal�ontologie
Monatsheft. 1996(10), 635-644.
Hutt, 1999. Neovenator salerii: A new theropod dinosaur from the Wealden
of the Isle of Wight: Its status and significance for theropod evolution. Masters
thesis. University of Portsmouth. 196 pp.
Hutt, 2001. Catalogue of Wealden Group Dinosauria in the Museum of Isle of Wight
Geology. In Martill and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological
Association. 411-422.
Martill and Naish, 2001. The geology of the Isle of Wight. In Martill and Naish
(eds.). Dinosaurs of the Isle of Wight. The Palaeontological Association. 25-43.
Naish, Hutt and Martill, 2001. Saurischian dinosaurs 2: Theropods. In Martill
and Naish (eds.). Dinosaurs of the Isle of Wight. The Palaeontological Association.
242-309.
Sweetman, 2004. The first record of velociraptorine dinosaurs (Saurischia, Theropoda)
from the Wealden (Early Cretaceous, Barremian) of Southern England. Cretaceous
Research. 25, 353-364.
Brusatte, Benson and Hutt, 2008. The osteology of Neovenator salerii
(Dinosauria: Theropoda) from the Wealden (Barremian) of the Isle ofWight. Monograph
of the Palaeontographical Society. 162(631), 1-166.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Barker, Dyke, Naish, Newham and Katsamenis, 2015. Complex neurovascular network
in the rostrum of Neovenator salerii. SVPCA 2015 abstracts, 78.
Barker, Naish, Newham, Katsamenis and Dyke, 2017. Complex neuroanatomy in the
rostrum of the Isle of Wight theropod Neovenator salerii. Scientific
Reports. 7: 3749, 1-8.
Ulughbegsaurus Tanaka, Anvarov, Zelenitsky, Ahmedshaev and Kobayashi, 2021
U. uzbekistanensis Tanaka, Anvarov, Zelenitsky, Ahmedshaev and Kobayashi, 2021
Turonian, Late Cretaceous
Dzharakuduk, Bissekty Formation, Uzbekistan
Holotype- (UzSGM 11-01-02) (~7.5-8 m) partial maxilla
Paratypes- (CCMGE 600/12457) maxillary fragment
(ZIN PH 357/16) maxillary fragment
Diagnosis- (after Tanaka et
al., 2021) series of shallow, oval depressions on the lateral surface
along the ventral edge of the maxilla; tubercles along rim of
antorbital fossa; vertically oriented ridges on lateral surface of the
maxilla; large foramina at dorsal edge of paradental plates in maxilla.
Comments- Tanaka et al. (2020)
first published about this specimen in an abstract, then named it the
following year. Nessov (1995) referred the maxilla CCMGE
600/12457 to Alectrosaurus,
but it was reidentified as dromaeosaurid by Averianov and Sues (2012) and described
as cf. Itemirus by Sues and Averianov (2014). Tanaka et al. (2021) referred it to Ulughbegsaurus along with maxillary fragment ZIN PH 357/16 that was described as cf. Itemirus
by Sues and Averianov. Tanaka et al. (2021) also suggested "it is
possible that some of the large tyrannosauroid teeth reported from
Dzharakuduk of the Bissekty Formation (with CBLs up to 26.6 mm and CBR
to 0.41-0.70) could belong to carcharodontosaurians." Tanaka et
al. (2020) entered it into Carrano's Tetanurae analysis where it
emerged in a
polytomy with megaraptorans and Neovenator, while in the 2021 publication they also added it to Novas' tetanurine analysis and recovered in a polytomy with Neovenator, Siamraptor, Concavenator and Eocarcharia. Compared to Neovenator and Siamraptor, Ulughbegsaurus
is more similar to the former in the tall antorbital fossa below the
antorbital fenestra and less transversely compressed alveoli, but more
like Siamraptor in the dorsoventrally narrow exposure of the lateral alveolar wall in medial view.
References-
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust. 1-156.
Averianov and Sues, 2012. Skeletal remains of Tyrannosauroidea (Dinosauria:
Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan.
34, 284-297.
Sues and Averianov, 2014. Dromaeosauridae (Dinosauria: Theropoda) from the Bissekty
Formation (Upper Cretaceous: Turonian) of Uzbekistan and the phylogenetic position
of Itemirus medullaris Kurzanov, 1976. Cretaceous Research. 51, 225-240.
Tanaka, Anvarov,
Ahmedshaev and Kobayashi, 2020. A large neovenatorid (Dinosauria:
Theropoda) from the Upper Cretaceous Bissekty Formation (Turonian),
Uzbekistan. The Society of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 318.
Tanaka, Anvarov, Zelenitsky, Ahmedshaev and Kobayashi, 2021. A new
carcharodontosaurian theropod dinosaur occupies apex predator niche in
the early Late Cretaceous of Uzbekistan. Royal Society Open Science. 8:
210923.
Carcharodontosauridae sensu Benson, Carrano and Brusatte, 2010
Definition- (Carcharodontosaurus saharicus <- Neovenator salerii,
Allosaurus fragilis, Sinraptor dongi)
unnamed carcharodontosaurid (Gasca, Canudo and Moreno-Azanza, 2014)
Early Barremian, Early Cretaceous
Mirambel Formation, Spain
Material- (MPZ 2014/235) distal femur
Reference- Gasca, Canudo and Moreno-Azanza, 2014. A large-bodied theropod
(Tetanurae: Carcharodontosauria) from the Mirambel Formation (Barremian) of
Spain. Neues Jahrbuch f�r Geologie und Pal�ontologie Abhandlungen.273(1),
13-23.
unnamed carcharodontosaurid (Mo, Huang, Xie and Buffetaut, 2014)
Aptian?, Early Cretaceous
Xinlong Formation, Guangxi, China
Material- (NHMG 10858) lateral tooth (71x37x17 mm)
Reference- Mo, Huang, Xie and Buffetaut, 2014. A megatheropod tooth from
the Early Cretaceous of Fusui, Guangxi, Southern China. Acta Geologica Sinica
(English Edition). 88(1), 6-12.
"Osteoporosia" Molina-Perez and Larramendi, 2019
"O. gigantea" Molina-Perez and Larramendi, 2019
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Material- (JP Cr340) partial posterior dorsal neural arch
Comments- This was discovered in 2009 and briefly described online by
Singer (2015), on the website of a theme park chain based on a specimen collected
and owned by the chain. He assigns this unofficial taxon to Carnosauria, and
indeed it strongly resembles the posterior dorsal of Mapusaurus. Thus
it may belong to the contemporaneous Carcharodontosaurus or Saurionops,
but cannot be compared to either. Molina-Perez and Larramendi (2019)
published the name as a nomen nudum, stating ""Osteoporosia gigantea"
is the informal name of a supposed gigantic megaraptorid that was 15 m
long, but it seems to have been a significantly smaller
carcharodontosaurid."
References- Singer, 2015 online. JuraPark na tropie nowych dinozaurow
z Maroka. https://web.archive.org/web/20151206224352/https://jurapark.pl/jurapark-na-tropie-nowych-dinozaurow-z-maroka/
Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288 pp.
unnamed clade (Altispinax dunkeri + Concavenator corcovatus
+ Eocarcharia dinops + Sauroniops pachytholus)
Diagnosis- (after Carrano et al., 2012) squared anterior margin of antorbital
fossa (unknown in Altispinax and Sauroniops)
(after Cau et al., 2012) prefrontal facet of the frontal restricted to the anterior
end of the lateral surface (unknown in Altispinax and Concavenator);
wide nasal-frontal contact reaching the posteromedial end of the nasal process
of the frontal (unknown in Altispinax and Concavenator); preorbital
facet that is deepest in the anterior end of the facet (unknown in Altispinax
and Concavenator).
(suggested) accessory centrodiapophyseal lamina on the transverse processes
of the posterior dorsal vertebrae (unknown in Eocarcharia and Sauroniops);
eleventh and twelfth dorsal neural spines highly elongate (five times the height
of the centra), while ninth dorsal neural spine is short (unknown in Eocarcharia
and Sauroniops); firm contact between apexes of eleventh and twelfth
dorsal neural spines (unknown in Eocarcharia and Sauroniops).
Altispinax Huene, 1923
= Becklespinax Olshevsky, 1991
A. dunkeri Huene, 1923
= Acrocanthasaurus altispinax Paul 1988
= Becklespinax altispinax (Paul, 1988) Olshevsky, 1991
= Altispinax "lydekkerhueneorum" Pickering, 1984 vide Pickering,
1995
= Altispinax altispinax (Paul, 1988) Rauhut, 2000
Valanginian, Early Cretaceous
Hastings Beds Group, England
Holotype- (NHMUK R1828) (~5 m) tenth dorsal vertebra, eleventh dorsal
vertebra, twelfth dorsal vertebra, two vertebrae (lost), two ribs (lost)
Diagnosis- (after Naish, 2011) posterior dorsal spines transversely expanded
at tips; large. rectangular hyposphene in posterior dorsals (unknown in Concavenator).
(after Cuesta et al., 2013) (compared to Concavenator) less ventrally
constricted centra; anteriorly angled posterior dorsal neural spines; posterior
dorsal neural spines distally expanded anteroposteriorly; posterior dorsal neural
spines transversely broader.
Other diagnoses- Olshevsky (1991) listed "firm contact between the
apexes of the neural spines of vertebrae #9 and 10 (and, presumably, later vertebrae
in the series)", but this is also present in Concavenator (though
the vertebrae are 11 and 12 in the latter genus and presumably Becklespinax
too).
Comments- This taxon has a complicated nomenclatural history. Altispinax
was initially proposed by Huene (1923) for "the species described as M.
dunkeri by Lydekker (Dames)" ... "distinguished from Megalosaurus
by its enormously high neural spines in the dorsal region" (referencing
three high-spined dorsal vertebrae catalogued as NHMUK R1828), which was misunderstood
by future authors as proposing a new genus for Dames' (1884) Megalosaurus
dunkeri. As M. dunkeri's holotype is an incomparable tooth, Stovall
and Langston (1950) tried to designate NHMUK R1828 (as BMNH R1828) as the type of "Altispinax
dunkeri (Dames)", but Dames' species already had a holotype which could
not be replaced without an ICZN petition. Paul (1988) solved this by creating
the new species altispinax for the vertebrae. Paul placed altispinax
in Acrocanthosaurus, but Olshevsky (1991) disagreed and created the new
genus Becklespinax for these vertebrae. Both of these options left Dames'
tooth as Altispinax dunkeri, which has been the consensus for over a
decade. Rauhut (2000; published as 2003) used a quote from Huene (1932) to justify
that author's intent to base Altispinax on the vertebrae, and so created
the combination Altispinax altispinax for them. Finally, Maisch (2016)
recognized that Huene's initial quote had been misunderstood, proposing Altispinax
for "the species described as M. dunkeri by Lydekker (Dames)",
not the species M. dunkeri (Dames) itself. Utilizing ICZN Article 11.10
("If an author employs a specific or subspecific name for the type species
of a new nominal genus-group taxon, but deliberately in the sense of a previous
misidentification of it, then the author's employment of the name is deemed
to denote a new nominal species and the specific name is available with its
own author and date as though it were newly proposed in combination with the
new genus-group name"), Maisch determined the species dunkeri Huene,
1923 is deemed to be different than dunkeri Dames, 1884. Thus Altispinax
dunkeri Huene, 1923 is a new taxon different from Megalosaurus dunkeri
Dames, 1884. Note Kuhn (1939) was the first to designate dunkeri as the
type species of Altispinax (Huene also initially referred Megalosaurus
oweni to it, now the type species of Valdoraptor).
Pickering (1995) attempted to make NHMUK R1828 (as BMNH R1828) the lectotype of Altispinax
"lydekkerhueneorum", which included as paratypes the holotype of Valdoraptor
oweni and several additional specimens (NHMUK R604, 604a-b, 604d). However,
the species altispinax, dunkeri and oweni have priority (Pickering
incorrectly considered them nomina rejecta, which cannot occur without an ICZN
decision), and there is no evidence these specimens are conspecific. This makes
Pickering's species (which is a nomen nudum in any case) an objective junior
synonym of Altispinax dunkeri.
Note abundant additional material was referred to Megalosaurus dunkeri
by Lydekker (1888), then Altispinax dunkeri by Huene (1926), but has
not been described in detail so cannot be compared to the holotypes of either.
The contemporaneous Hastings Beds specimens are listed below, while the Weald
Clay (Belgium, England) and Lower Greensand (England) material is listed as Averostra here.
The holotype dorsals were first referred to Megalosaurus bucklandii by
Owen (1855, 1857), before the nomenclatural saga of Altispinax described
above began. After decades of being assigned largely by default to Megalosauridae,
or to Spinosauridae due to the neural spine elongation, NHMUK R1828 has more
recently been assigned to Allosauridae as Acrocanthosaurus (Paul, 1988),
Eustreptospondylidae based on similarity to Piatnitzkysaurus (Olshevsky,
1991), Allosauroidea (Naish, 1999) or Carcharodontosauridae (Naish, 2011; his
Carcharodontosauria). The latter is supported by its similarity to the recently
discovered Concavenator. However, Carrano et al. (2012) believe the anterior
dorsal neural spine is broken, lessening the resemblence. While Carrano et al.
state that within Tetanurae it can only be excluded from Spinosauridae and Carcharodontosauridae,
when entered conservatively into their matrix it emerges as a carnosaur and
most often sister to Concavenator and Eocarcharia. The discrepancy
is due to the authors stating Altispinax is unlike carcharodontosaurids
in lacking posterior dorsal pleurocoels, but Concavenator also lacks
these.
References- Owen, 1855. Monograph on the Fossil Reptilia of the Wealden
and Purbeck Formations. Part II. Dinosauria (Iguanodon) (Wealden). Palaeontographical
Society Monographs. 8, 1-54.
Owen, 1857. Monograph on the Fossil Reptilia of the Wealden and Purbeck Formations.
Part III. Dinosauria (Megalosaurus) (Wealden). Palaeontographical Society
Monographs. 9, 1-26.
Dames, 1884. Vorlegung eines Zahnes von Megalosaurus aus dem Wealden
des Deisters. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu
Berlin. 1884, 186-188.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of
the Geological Society of America. 34, 449-458.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Kuhn, 1939. Saurischia. In Quenstedt (ed.). Fossilium Catalogus, I: Animalia,
pars 87, 124 pp.
Stovall and Langston, 1950. Acrocanthosaurus atokensis, a new genus and
species of Lower Cretaceous Theropoda from Oklahoma. The American Midland Naturalist.
43(3), 696-728.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry,
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Naish, 1999. Studies on Wealden Group theropods - an investigation into the
historical taxonomy and phylogenetic affinities of new and previously neglected
specimens. MPhil thesis, University of Portsmouth. [pp]
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Cuesta, Ortega and Sanz, 2013. Solving the synonymy issue in Concavenator
corcovatus and Becklespinax altispinax, two distinct theropods from
the Lower Cretaceous of Europe. Journal of Vertebrate Paleontology. Program
and Abstracts 2013, 110.
Maisch, 2016. The nomenclatural status of the carnivorous dinosaur genus Altispinax
v. Huene, 1923 (Saurischia, Theropoda) from the Lower Cretaceous of England.
Neues Jahrbuch f�r Geologie und Pal�ontologie - Abhandlungen, 280(2),
215-219.
A? sp. indet. (Benton and Spencer, 1995)
Valanginian, Early Cretaceous
Hastings Beds, England
Material- (NHMUK R1954) teeth, limb elements
teeth, vertebrae
Comments- This material was referred to Altispinax dunkeri without
justification by Benton and Spencer (1995).
Reference- Benton and Spencer, 1995. Fossil Reptiles of Great Britain.
Chapman & Hall, London. 386 pp.
A? sp. indet. (Lydekker, 1888)
Middle Valanginian, Early Cretaceous
Wadhurst Clay of the Hastings Beds, England
Material- (NHMUK 39213) tooth (Lydekker, 1888)
(NHMUK R604; intended paratype of Altispinax "lydekkerhueneorum")
two teeth (Lydekker, 1888)
(NHMUK R604a; intended paratype of Altispinax "lydekkerhueneorum")
incomplete anterior dorsal vertebra (Lydekker, 1888)
(NHMUK R604b; intended paratype of Altispinax "lydekkerhueneorum")
incomplete scapula (Lydekker, 1888)
(NHMUK R604c; intended paratype of Altispinax "lydekkerhueneorum")
partial tibia (Lydekker, 1888)
....(NHMUK R604d; intended paratype of Altispinax "lydekkerhueneorum")
metatarsal IV (245 mm) (Lydekker, 1888)
(NHMUK R641) three teeth (Lydekker, 1888)
(NHMUK R1525) metatarsal II (Huene, 1926)
(NHMUK R1525a) incomplete mid caudal vertebra (Huene, 1926)
Comments- Most of this material was referred to Megalosaurus dunkeri
by Lydekker (1888), then Altispinax dunkeri by Huene (1926), but has
not been described in detail so cannot be compared to the holotypes of either.
They may belong to Baryonyx, Altispinax, Valdoraptor, Neovenator, Calamosaurus,
Eotyrannus or another Wealden theropod. NHMUK R604a, 604c-d and 1525 were
referred to Valdoraptor (then Megalosaurus) oweni by Lydekker
(1890), but this has not been accepted by recent workers. Pickering (1995) intended
to use the 604 specimens as paratypes of his proposed species Altispinax
"lydekkerhueneorum", but this is a nomen nudum and objective junior
synonym of Altispinax dunkeri (see entry).
References- Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia
in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing
the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia,
and Proterosauria. British Museum of Natural History, London. 309 pp.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous Formations,
principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry,
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Concavenator Ortega, Escaso
and Sanz, 2010
C. corcovatus Ortega, Escaso and Sanz, 2010
Late Barremian, Early Cretaceous
Calizas de La Huerguina Formation, Spain
Holotype- (MCCM-LH 6666) incomplete skull, partial mandible, ten cervical
vertebrae, cervical ribs, thirteen dorsal vertebrae, dorsal ribs, gastralia,
five sacral vertebrae, thirty caudal vertebrae, twenty-six chevrons, scapulae,
coracoid, humeri, radius, ulna, metacarpal I, phalanx I-1, metacarpal II, phalanx
II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-3, manual
ungual III, ilium, pubes, ischium, femora (one incomplete), tibiae (one incomplete),
incomplete fibula, astragalus, calcaneum, two distal tarsals, metatarsals II,
phalanges II-1 (one proximal), phalanx II-2 fragment, metatarsals III, phalanx
III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx
IV-1, partial phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal
V, pedal claw sheaths, scale impressions, smaller theropod vertebrae
Diagnosis- (after Ortega et al., 2010) four nasal recesses, three of
them connected (unknown in Altispinax); alarge, rounded, thickened postorbital
brow occupying one-third of the orbit (unknown in Altispinax); relatively
high, anteriorly directed neurapophyses of second and third caudal vertebrae
(unknown in Altispinax); small, thorn-like posterior process at base
of each neurapophysis in proximal caudal vertebrae (unknown in Altispinax).
(after Carrano et al., 2012) differs from Altispinax in- posterior dorsal
centra bear large lateral fossae (although no foramina); posterior dorsal neural
spines have less curved anterior and posterior margins at their bases; apices
of the tallest dorsal neural spines are anteroposteriorly narrow, curving towards
a single apex formed from multiple spines.
(proposed) tenth dorsal neural spine one third as tall as eleventh dorsal neural
spine; eleventh dorsal neural spine strongly tapered distally in lateral view;
eleventh and twelfth dorsal neural spines appressed except at base.
Other diagnoses- Ortega et al. (2010) also included "tall neurapophyses
of the eleventh and twelfth dorsal vertebrae (five times the height of the centra)"
in their diagnosis, but this is also true in Altispinax.
Comments- Ortega et al. (2010) identified a line along the lateral side
of Concavenator's ulna as possessing quill knobs, which are otherwise
unknown in non-paravian taxa. However, both Mortimer (online, 2010) and Naish
(online, 2010) independantly regard the structure as an intermuscular line instead,
with the former identifying it more precisely as the line between the extensor
carpi ulnaris and the flexor ulnaris. This is due the more anterior position
of the line, which coincides with the intermuscular boundary on Alligator
and non-maniraptoran theropods, but not with quill knobs in paravians.
References- Mortimer, online 2010. http://theropoddatabase.blogspot.com/2010/09/concavenator-part-ii-becklespinax.html
Naish, online 2010. https://web.archive.org/web/20100911131144/http://scienceblogs.com/tetrapodzoology/2010/09/concavenator_incredible_allosauroid.php
Ortega, Escaso and Sanz, 2010. A bizarre, humped Carcharodontosauria (Theropoda)
from the Lower Cretaceous of Spain. Nature. 467, 203-206.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Cuesta, Ortega and Sanz, 2013. Solving the synonymy issue in Concavenator
corcovatus and Becklespinax altispinax, two distinct theropods from
the Lower Cretaceous of Europe. Journal of Vertebrate Paleontology. Program
and Abstracts 2013, 110.
Cuesta, Diaz-Martinez, Ortega and Sanz, 2014. Phylogenetic implications of the
bird-like podotheca of Concavenator corcovatus (Theropoda, Carcharodontosauridae).
Journal of Vertebrate Paleontology. Program and Abstracts 2014, 114-115.
Cuesta, Ortega and Sanz, 2015. Ulnar bumps of Concavenator: Quill knobs
or muscular scars? Myological reconstruction of the forelimb of Concavenator
corcovatus (Lower Cretaceous, Las Hoyas, Spain). Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 111-112.
Eocarcharia Sereno and Brusatte,
2008
E. dinops Sereno and Brusatte, 2008
Aptian-Albian, Early Cretaceous
Elrhaz Formation of the Tegama Group, Niger
Holotype- (MNN GAD2) (~6-8 m) postorbital (163 mm tall)
Paratypes- (MNN GAD3) postorbital
(MNN GAD4) partial postorbital
(MNN GAD5) partial postorbital
(MNN GAD6) partial postorbital
(MNN GAD7) incomplete maxilla (528 mm)
(MNN GAD8) maxillary fragment
(MNN GAD9) maxillary fragment
(MNN GAD10) prefrontal (55 mm), frontal (102 mm)
(MNN GAD11) frontoparietal, orbitosphenoid fragments
(MNN GAD12) three teeth
(MNN GAD13) tooth fragment
(MNN GAD14) tooth (48 mm)
Referred- ?(MNN GAD15) tooth (Sereno and Brusatte, 2008)
? teeth (Sereno et al., 1994)
? vertebrae, pelvic elements including pubis (Sereno, unpublished)
Diagnosis- (after Sereno and Brusatte, 2008) enlarged subtriangular laterally
exposed promaxillary fenestra larger in size than the maxillary fenestra; circular
accessory pneumatic fenestra on the posterodorsal ramus of the maxilla; dorsoventral
expansion of the antorbital fossa ventral to the promaxillary and maxillary
fenestrae; postorbital brow accentuated by a finely textured boss, positioned
above the posterodorsal corner of the orbit; postorbital medial process with
a plate-shaped projection fitted to an articular slot on the frontal; postorbital
articulation for the jugal that includes a narrow laterally-facing facet; enlarged
prefrontal lacking the ventral process with subquadrate exposure on the dorsal
skull roof and within the orbit (limiting the anterior ramus of the frontal
to the roof over the olfactory bulbs); low protuberance on the frontoparietal
suture.
Comments- Sereno et al. (1994) noted mid-sized carcharodontosaurid teeth
from the Elrhaz Formation. Sereno later mentioned carcharodontosaurid remains
from Gadoufaoua in an online update to his 2000 Project Exploration dig in Niger.
These included teeth, jaw bones and a postorbital (which are elements described
for Eocarcharia), but also vertebrae and pelvic elements. A pubis was
photographed in situ. These may be elements later referred to Kryptops,
as no Eocarcharia postcrania were described in Sereno and Brusatte (2008).
Kryptops' pelvis and dorsals do look somewhat carnosaurian, but the in
situ pubis does not appear to be the Kryptops specimen.
Novas et al. (2013) believed there was not enough evidence to refer the maxillary
and dental material to Eocarcharia, but given the presence of a single
type of carcharodontosaurid skull roof in the formation, it seems probable the
jaw bones belong to the same taxon.
References- 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.
Sereno, online 2000. https://web.archive.org/web/20020417224123/http://www.projectexploration.org/niger2000/10_03_2000_2.htm
Brusatte and Sereno, 2006. Basal abelisaurid and carcharodontosaurid theropods
from the Elrhaz Formation (Aptian-Albian) of Niger. Journal of Vertebrate Paleontology.
27(3), 46A.
Sereno and Brusatte, 2008. Basal abelisaurid and carcharodontosaurid theropods
from the Lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica Polonica.
53(1), 15-46.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Lajasvenator Coria, Currie, Ortega and Baiano, 2020
= "Lajasvenator" Coria, Currie, Ortega and Baiano, 2019
L. ascheriae Coria, Currie, Ortega and Baiano, 2020
= "Lajasvenator ascheriae" Coria, Currie, Ortega and Baiano, 2019
Late Valanginian, Early Cretaceous
Mulichinco Formation, Neuquen, Argentina
Holotype- (MLL-PV-005) (adult)
premaxillae (one incomplete), maxillae (one partial, one fragmentary),
anterior vomer, posterior dentary, partial splenial, partial seventh
cervical vertebra (102 mm), eighth cervical vertebra (90 mm), ninth
cervical vertebra (103 mm), tenth cervical vertebra (92 mm), cervical
ribs (260, 220 mm), incomplete first dorsal vertebra, posterior fifth
dorsal centrum, sixth dorsal centrum (71 mm), seventh dorsal centrum
(71.4 mm), partial eighth dorsal vertebra (72.6 mm), partial ninth
dorsal vertebra, partial tenth dorsal vertebra (68.7 mm), partial
eleventh dorsal vertebra (74.2 mm), partial twelfth dorsal vertebra
(71.3 mm), anterior thirteenth dorsal centrum, five incomplete
posterior dorsal ribs (350, 250, 210, 170 mm), several dorsal rib
fragments, four ?gastralial fragments, incomplete fused second to
fourth sacral vertebrae (s3 68 mm), three mid caudal vertebrae (66, 67
mm), mid caudal vertebral fragment, fragmentary ilium, proximal pubis,
proximal ischial fragment
Paratype- (MLL-PV-007)
quadratojugal, anterior dentaries, four cervical or anterior dorsal
transverse processes, proximal seventh cervical rib, proximal anterior
dorsal rib, radiale or distal tarsal, distal metatarsal, fragments
Diagnosis- (after Coria et al.,
2019) anterior projection on cervical prezygapophyses; lip-like crest
on lateral surface of cervical postzygapophyses; cervical ribs with
bilobed anterior process.
Comments- The holotype was
discovered in 2010 and the paratype in 2012. Coria et al. (2010)
initially wrote "certain anatomical features of the collected specimens
suggest abelisaur affinities in the theropods", but Coria et al. (2019)
later described it as a carnosaur. Coria et al. (2019) named and
described
the specimen as Lajasvenator ascheriae
in a journal pre-proof posted November 25 2019, but this was electronic
and had no mention of ZooBank, so it was a nomen nudum (ICZN Article
8.5.3. states names published electronically must "be registered in the
Official Register of Zoological Nomenclature (ZooBank) (see Article
78.2.4) and contain evidence in the work itself that such registration
has occurred") until July 2020.
Coria et al. (2019) added it to Carrano et al.'s tetanurine analysis and found it to form a clade in Carcharodontosauridae with Eocarcharia and Concavenator.
References- Coria, Currie,
Koppelhus, Braun and Cerda, 2010. First record of a Valanginian (Early
Cretaceous) dinosaur association from South America. Journal of
Vertebrate Paleontology 30: 75A.
Coria, Currie, Ortega and Baiano, 2020 (online 2019). An Early Cretaceous,
medium-sized carcharodontosaurid theropod (Dinosauria, Saurischia) from
the Mulichinco Formation (upper Valanginian), Neuqu�n Province,
Patagonia, Argentina. Cretaceous Research. 111, 104319.
Sauroniops Cau, Dalla Vecchia
and Fabbri, 2013
S. pachytholus Cau, Dalla Vecchia and Fabbri, 2013
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Holotype- (MPM 2594) frontal (186 mm)
Diagnosis- (after Cau et al., 2013) dorsoventrally thickened frontal
(with the depth of the body ranging between 28%, along the medial suture, and
38%, at the level of the anteromedial margin of the supratemporal fossa, of
bone length); nasal processes of the frontal with a transversely convex dorsal
surface and completely separated medially by the posteromedial processes of
nasal extended along 40% of the frontal length and reaching the frontal body;
thick dome-like eminence in the anterolateral corner of the dorsal surface of
the frontal at the level of the prefrontal-lacrimal articulations; anterolateral
margin of the lateral surface of the frontal with a narrow vertical lamina separating
the prefrontal facet from an elliptical fossa in the lacrimal facet; prefrontal
facet of the frontal trapezoidal, barely visible in ventral view, mostly restricted
to the anterodorsal margin of the lateral surface of the bone, and not participating
in the orbital fossa; frontal with interdigitate suture for prefrontal restricted
to the anteroventral corner of the facet, formed by a low shelf running along
the posterolateral margin of the nasal process and a small finger-like projection;
hypertrophied, D-shaped lacrimal facet of the frontal bordering the whole posterolateral
exposure of the prefrontal facet and with maximum depth that is four times the
depth of the anterior half of the postorbital facet; dorsal surface of the frontal
anterior to the anteromedial margin of supratemporal fossa raised and facing
anterodorsally, describing with the dorsal dome a posteromedially-anterolaterally
directed saddle-shaped concavity, and confluent with a series of low rounded
rugosities placed posteriorly to the nasal facet.
Comments- Cau et al. (2012, 2013) found this specimen to be most similar
to Eocarcharia.
Ibrahim et al. (2020) propose Sauroniops is an immature Carcharodontosaurus saharicus,
but this is based on misconceptions. First, they state it is
"approximately 60% the size of the frontal in the original holotype and
neotype of C. saharicus" based on the length posterior to the lateral nasal contact, but the width (~153 mm) is actually greater than the Carcharodontosaurus neotype (~124 mm). This shows it is not a smaller specimen and the broadeer proportions are more like Eocarcharia. Ibrahim et al. then claim "a prefrontal articular facet on the frontal in S. pachytholus is said to be absent in C. saharicus", but Cau et al. state in Carcharodontosaurus
"the prefrontal facet is a rounded pit placed anterodorsally to the
orbital margin and in the posterodorsal corner to the lacrimal facet,
thus approaching the postorbital facet." Ibrahim et al. say
"other immature skull elements of C. saharicus
found in the Kem Kem Group, such as a postorbital, show less prominent
development of brow ornamentation as one would anticipate in the course
of growth" as if to defend an immature age for Sauroniops, but it's Sauroniops that has the "unique presence of dorsal ornamentations and eminence" compared to Carcharodontosaurus. Ibrahim et al. further misunderstand Cau et al.'s wording, as the diagnosis states it differs from Carcharodontosaurus
"in lacking a dorsomedially inclined anterior half of the dorsal
surface of the frontal, a deeply invaginated anterior margin of the
supratemporal fossa, prominent frontal shelves overhanging the
supratemporal fossa, and an extensively ossified interorbital septum",
whereas Ibrahim think they mean Sauroniops
has frontal shelves and an ossified interorbital septum. So these
stand as valid differences between the two taxa. Cau et al. use
the elongate posteromedial nasal process to distinguish Sauroniops from Carcharodontosaurus, so it's not true that "C. saharicus
shows a similar W-shaped nasal-frontal suture." The only
objection of Ibrahim et al. that is correct is that "only the central
portion of the supratemporal fossa is preserved in S. pachytholus", so that it cannot be said to differ from C. saharicus "in showing a distinct anterolateral corner instead of a more gently curved anterolateral margin in dorsal view." Sauroniops is retained as valid here.
References- Cau, Dalla Vecchia and Fabbri, 2012. Evidence of a new carcharodontosaurid
from the Upper Cretaceous of Morocco. Acta Palaeontologica Polonica. 57(3),
661-665.
Cau, Dalla Vecchia and Fabbri, 2013. A thick-skulled theropod (Dinosauria, Saurischia)
from the Upper Cretaceous of Morocco with implications for carcharodontosaurid
cranial evolution. Cretaceous Research. 40, 251-260.
Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020. Geology and paleontology of the Upper
Cretaceous Kem Kem Group of eastern Morocco. ZooKeys. 928, 1-216.
Acrocanthosaurus Stovall
and Langston, 1950
= "Acrocanthus" Langston, 1947
Comments- Goodwin et al. (1999) referred teeth from the Mugher Mudstone
of Ethiopia to cf. Acrocanthosaurus sp., but these can only be identified
as Tetanurae indet..
Reference- Goodwin, Clemens, Hutchison, Wood, Zavada, Kemp, Duffin and
Schaff, 1999. Mesozoic continental vertebrates with associated palynostratigraphic
dates from the northwestern Ethiopian plateau. Journal of Vertebrate Paleontology.
19(4), 728-741.
A. atokensis Stovall and Langston, 1950
= "Acracanthus atokaensis" Langston, 1947
Late Aptian-Middle Albian, Early Cretaceous
Antlers Formation, Oklahoma, US
Holotype- (MOU 8-0-S9 or OMNH 10146) (9.9 m) lacrimal, partial jugal,
partial postorbital, incomplete squamosal, frontals, parietals, braincase, ectopterygoid,
articular, fragment of surangular, fragment of angular, atlantal intercentrum,
axial fragment, partial third cervical vertebra (96 mm), incomplete fourth cervical
vertebra (98 mm), incomplete fifth cervical vertebra (123 mm), sixth cervical
neural arch, partial seventh cervical centrum (153 mm), incomplete eighth cervical
vertebra (158 mm), ninth cervical vertebra (168 mm), incomplete tenth cervical
vertebra (153 mm), three incomplete cervical ribs, fifth dorsal vertebra (107
mm), sixth dorsal vertebra (110 mm), seventh dorsal vertebra, twelfth dorsal
centrum (128 mm), thirteenth dorsal centrum (125 mm), eight dorsal neural spines,
five posterior dorsal ribs, gastralium, two caudal centra, two proximal chevrons,
coracoid, pubes (~838 mm), ischium (~621 mm), distal femur (~950 mm), tibia
(~865 mm), fibulae (801 mm), astragalus, metatarsal III (416 mm)
Paratype- (MOU 8-0-S8 or OMNH 10147) (11.29 m) two dorsal centra, four
dorsal neural spines, eight posterior dorsal ribs, first caudal vertebra, second
caudal vertebra (128 mm), third caudal vertebra (138 mm), fourth caudal vertebra
(140 mm), ninth caudal vertebra (149 mm), tenth caudal vertebra (146 mm), eleventh
caudal vertebra (141 mm), twelfth caudal vertebra (140 mm), eighteenth caudal
vertebra, nineteenth caudal vertebra (131 mm), twentieth caudal vertebra (134
mm), twenty-first caudal vertebra (135 mm), twenty-second caudal vertebra, twenty-third
caudal vertebra (124 mm), proximal chevron, pubes, proximal femur (~950 mm),
fragmentary tibia (~958 mm), metatarsal II (416 mm), metatarsal III (445 mm),
phalanx III-1 (145 mm)
Referred- (NCSM 14345; OMNH 10168) (11.5 m, adult) skull (1.29 m), mandibles
(1.315 m), presacral vertebral fragments, cervical rib, several partial dorsal
ribs, gastralia fragments, over fourteen caudal vertebrae (120-160 mm), six
chevrons, scapula (970 mm), coracoid (360 mm), humerus (370 mm), radius (220
mm), ulna (255 mm), radiale, intermedium, semilunate carpal, metacarpal I (62 mm),
phalanx I-1 (111 mm), proximal manual ungual I, metacarpal II (116 mm), phalanx
II-1 (101 mm), proximal manual ungual II (124, 144 mm), metacarpal III (89 mm),
phalanx III-1 (50 mm), phalanx III-2 (42 mm), incomplete femur (~1.277 m), incomplete
tibia, partial astragalus, calcaneum, metatarsal I (111 mm), phalanx I-1 (70
mm), pedal ungual I, metatarsal II (410 mm), phalanx II-1 (55 mm), phalanx II-2
(122 mm), partial metatarsal III (~439 mm), phalanx III-1 (160 mm), phalanx
III-2 (115 mm), partial metatarsal IV, phalanx IV-1 (85 mm), phalanx IV-2 (70
mm), phalanx IV-3 (58 mm), phalanx IV-4, pedal ungual IV, metatarsal V (200
mm) (Currie and Carpenter, 2000)
?(OMNH 3031) four gastralia (paleofile.com)
(OMNH 51788) tooth (Lipka, 1998)
? teeth (Nydam et al., 1997)
Middle-Late Aptian
Arundel Formation, Maryland, US
(USNM 497718) tooth (Lipka, 1998)
(USNM 497722) tooth (Lipka, 1998)
(USNM 497723) tooth (Lipka, 1998)
(USNM 497724) tooth (Lipka, 1998)
(USNM 497725) tooth (Lipka, 1998)
(USNM 497726) tooth (Lipka, 1998)
Early Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
(CEU 5107) tooth (Kirkland et al., 1997)
Albian-Cenomanian, Early-Late Cretaceous
Turney Ranch Formation, Arizona, US
(ASDM coll.) tooth (Ratkevich, 1997)
Early Albian, Early Cretaceous
Paluxy Formation of the Trinity Group, Texas, US
(FMNH PR 965) tooth (72.8 mm) (Langston, 1974)
Aptian-Middle Albian, Early Cretaceous
Trinity Group, Texas
?(SMU 62271) teeth (Thurmond, 1974)
postorbital (Langston, 1974)
? femur (Langston, 1974)
(FMNH 2-51#1) juvenile premaxillary tooth fragment, seven lateral tooth fragments
(Gallup, 1975)
(FMNH 3-51#1) seven tooth fragments (Gallup, 1975)
(FMNH 8-52#1) premaxillary tooth fragment ( mm) (Gallup, 1975)
(FMNH 9s-52#1) tooth fragment (Gallup, 1975)
(FMNH 47-50) tooth (15.9 mm), tooth fragment (Gallup, 1975)
(FMNH 123-50) (juvenile) tooth (11.7 mm) (Gallup, 1975)
(FMNH 124-50) tooth (19 mm) (Gallup, 1975)
(FMNH 202-50) premaxillary tooth fragment, two lateral tooth fragments (Gallup,
1975)
(FMNH 209-50) tooth (97 mm) (Gallup, 1975)
(FMNH 212-50) tooth (47 mm) (Gallup, 1975)
(FMNH 239-50) two tooth fragments (Gallup, 1975)
(FMNH Turtle Gully) two tooth fragments (Gallup, 1975)
Aptian, Early Cretaceous
Twin Mountains Formation, Texas, US
(SMU 74646) (~9.8 m, 1.87 tons) incomplete jugal, ectopterygoid, palatine, partial surangular,
articular, partial prearticular, partial splenial, rostral tooth (84 mm tall,
31 by 19.5 mm wide), partial tooth, axis (101 mm), partial third cervical vertebra,
partial fourth cervical vertebra, partial fifth cervical vertebra (158 mm),
partial sixth cervical vertebra (180 mm), seventh cervical neural spine, eighth
cervical neural spine, ninth cervical neural spine, tenth cervical centrum,
two posterior cervical zygapophyseal assemblies, seven partial cervical ribs,
first dorsal centrum (295 mm), partial second dorsal vertebra (268 mm),partial
third dorsal vertebra, partial fourth dorsal vertebra, partial fifth dorsal
centrum, partial sixth dorsal vertebra, partial seventh dorsal vertebra, partial
eighth dorsal vertebra, incomplete ninth dorsal vertebra (135 mm), incomplete
tenth dorsal vetrtebra (144 mm), partial eleventh dorsal vertebra, partial twelfth
dorsal vertebra, partial thirteenth dorsal vertebra, ten dorsal neural spines,
nineteen partial dorsal ribs, dorsal rib fragments, gasteralia, partial first
sacral vertebra (170 mm), incomplete second sacral vertebra (160 mm), incomplete
third sacral vertebra (160 mm), partial fourth sacral vertebra, fifth sacral
fragment, two sacral neural spines, first caudal vertebra (117 mm), second caudal
vertebra (124 mm), fifth caudal vertebra (139 mm), sixth caudal vertebra (135
mm), eighth caudal vertebra (135 mm), fifteenth caudal vertebra (140.5 mm),
sixteenth caudal vertebra (150 mm), seventeenth caudal vertebra (142 mm), eighteenth
caudal vertebra (141 mm), ninteenth caudal vertebra (141 mm), twenty-second
caudal vertebra, twenty-eighth caudal vertebra (133 mm), twenty-ninth caudal
vertebra (131 mm), thirtieth caudal vertebra, proximal scapula, distal scapula,
incomplete pubes, ischia (844 mm), femora (1090 mm), distal metatarsal II (Harris,
1998)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Montana, US
?(YPM 5285) incomplete proximal caudal vertebra (103 mm) (Ostrom, 1970)
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Wyoming, US
(UM
20796) (4+ year old juvenile) dorsal centrum (100 mm), caudal neural
arch, incomplete pubes, femur (670 mm), proximal fibula, fragments
(Melstrom and D'Emic, 2011; described by D'Emic, Melstrom and Eddy,
2012)
(UM 20797) partial scapulocoracoid (D'Emic, Melstrom and Eddy, 2012)
?(YPM 5377) tooth (71x~26x? mm) (Ostrom, 1970)
Cenomanian, Late Cretaceous
Woodbine Formation, Texas, US
? teeth, limb fragments, manual ungual and/or pedal ungual (Main, Noto and
Scotese, 2011)
? teeth (Bennett et al., 2012)
Diagnosis- (after Carrano et al., 2012) absence of nasal extension of
antorbital fossa and associated pneumatopores; supraoccipital expanded parasagittally
into double boss posterior to nuchal crest; cervical vertebral neural spines
with triangular anterior processes that insert into fossae ventral to overhanging
processes on preceding neural spines; neural spines of presacral, sacral and
anterior caudal vertebrae more than 2.5 times taller than respective centrum
lengths; accessory process on lateral surface of caudal prezygapophysis
Comments- Langston (1947) originally called this taxon "Acrocanthus
atokaensis" in his thesis before describing it as Acrocanthosaurus
in 1950. The initial nomen nudum was published by Czaplewski et al. (1994).
Harris (1998) believes the tooth (UUVP 904) described by DeCourten (1991) does
not belong to Acrocanthosaurus because the serrations are too coarse.
However, Kirkland et al. (1997) mention cf. Acrocanthosaurus sp., which
is a tooth (CEU 5107) with finer serrations (Harris, 1998).
As stated by Currie and Carpenter (2000) "Parts of another specimen
(two posterior cervical or anterior dorsal centra, ischial fragment,
distal end of a femur), described in this paper, were collected by the
Oklahoma Museum of Natural History and were catalogued as OMNH 10168.
Most of the specimen, however, was collected by Cephis Hall & Sid
Love, who retrieved the skull, about two dozen vertebrae, ribs,
chevrons, most of the front limbs, pelvic fragments, parts of both
femora and tibiae, and most of the bones of the foot. ... The
unprepared specimen was eventually acquired by Allen Graffham of
Geological Enterprises, Inc., Ardmore, Oklahoma, who in turn arranged
for the Black Hills Institute in Hill City, South Dakota, to prepare
the specimen. The preparation was completed at the end of the summer of
1996, and the original skeleton went the following year to the North
Carolina State Museum of Natural Sciences as NCSM 14345. Carrano
(1998) includes measurements of this specimen as "Geol. Enterprises".
SMU 62271 were said to be similar to Allosaurus, so are provisionally
assigned to Acrocanthosaurus here based on provenance.
Senter and Robins (2005) note that NCSM 14345 does not include manual phalanges
II-2 or III-3, manual ungual III, or the tips of manual unguals I and II. These
were illustrated and described by Currie and Carpenter (2000) based on casts
made by BHI personnel.
Currie and Carpenter (2000) found Acrocanthosaurus to be an allosaurid,
but of their thirteen characters supporting this, at least four are primitive
(promaxillary and maxillary fenestrae; axial intercentrum subparallel to axis
ventral margin; paired anterior and posterior processes at base of chevrons;
pubic foramen present in distal pubis), three are also found in Giganotosaurus
(basioccipital participates in basal tubera; distal ends of paroccipital processes
below foramen magnum; internal carotid opening pneumatized), one in Carcharodontosaurus
(separation of trigeminal nerve branches complete), and five aren't known in
Carcharodontosaurus or Giganotosaurus (long basipterygoid processes;
reduced external mandibular fenestra; pronounced notch between acromion and
coracoid; sigmoidal humerus; metacarpal IV absent). So there are actually no
characters published by Currie and Carpenter that support placing Acrocanthosaurus
in the Allosauridae. Giganotosaurus and Carcharodontosaurus are
clearly more closely related to each other than Acrocanthosaurus is to
either, but there's no reason to believe the latter is not carcharodontosaurid.
When constrained as an allosaurid in Carrano et al.'s (2012) matrix, it takes
12 additional steps, showing this topology is indeed improbable.
References- Langston, 1947. A new genus and species of Cretaceous theropod
dinosaur from the Trinity of Atoka County, Oklahoma. MS Thesis, The University
of Oklahoma. 73 pp.
Stovall and Langston, 1950. Acrocanthosaurus atokensis, a new genus and
species of Lower Cretaceous Theropoda from Oklahoma. The American Midland Naturalist.
43(3), 696-728.
Langston, 1974. Nonmammalian Comanchean tetrapods. Geoscience and Man. 8, 77-102.
Thurmond, 1974. Lower vertebrate faunas of the Trinity division in North-Central
Texas. Geoscience and Man. 8, 103-129.
Gallup, 1975. Lower Cretaceous dinosaurs and associated vertebrates from north-central
Texas in the Field Museum of Natural History. MS thesis, University of Texas
at Austin. 159 pp.
DeCourten, 1991. The Long Walk quarry and tracksite: Unveiling the mysterious
Early Cretaceous of the Dinosaur Triangle region. In Averett (ed.). Guidebook
for dinosaur quarries and tracksites tour, Western Colorado and Eastern Utah.
Grand Junction: Grand Junction Geological Society. 19-25.
Czaplewski, Cifelli, and Langston, 1994. Catalog of type and figured fossil
vertebrates, Oklahoma Museum of Natural History. Oklahoma Geological Survey
Special Publication. 94, 1-35.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton,
1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau:
A key to understanding 35 million years of tectonics, sedimentology, evolution,
and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Nydam, Cifelli, Brinkman and Gardner, 1997. Preliminary report on the vertebrate
fauna of the Antlers Formation (Lower Cretaceous: Aptian-Albian) of Oklahoma.
Journal of Vertebrate Paleontology. 17(3), 67A.
Ratkevich, 1997. Dinosaur remains of southern Arizona. In Wolberg, Stump and
Rosenberg (eds.). Dinofest International. 213-221.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD thesis, The University of Chicago. 424
pp.
Harris, 1998. A reanalysis of Acrocanthosaurus atokensis, its phylogenetic
status, and paleobiogeographic implications, based on a new specimen from Texas.
New Mexico Museum of Natural History Bulletin. 13, 1-75.
Harris, 1998. Large, Early Cretaceous theropods in North America. In Lucas,
Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems.
New Mexico Museum of Natural History and Science Bulletin. 14, 225-228.
Lipka, 1998. The affinities of the enigmatic theropods of the Arundel Clay facies
(Aptian), Potomac Formation, Atlantic coastal plain of Maryland. In Lucas, Kirkland
and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico
Museum of Natural History and Science Bulletin. 14, 229-234.
Currie and Carpenter, 2000. A new specimen of Acrocanthosaurus atokensis
(Theropoda, Dinosauria) from the Lower Cretaceous Antlers Formation (Lower Cretaceous,
Aptian) of Oklahoma, USA. Geodiversitas. 22(2), 207-246.
Franzosa and Rowe, 2005. Cranial endocast of the Cretaceous theropod dinosaur
Acrocanthosaurus atokensis. Journal of Vertebrate Paleontology. 25(4),
859-864.
Senter and Robins, 2005. Range of motion in the forelimb of the theropod dinosaur
Acrocanthosaurus atokensis, and implications for predatory behaviour.
Zoological Journal. 266, 307-318.
Eddy, 2007. Results from a preliminary study of the bone histology of the Early
Cretaceous allosaurid Acrocanthosaurus atokensis. Journal of Vertebrate
Paleontology. 27(3), 70A.
Bates, 2008. Reconstructing the locomotor biology of Acrocanthosaurus atokensis
(Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 28(3), 48A.
Eddy, 2008. A re-analysis of the skull of Acrocanthosaurus atokensis
(NCSM 14345): Implications for allosauroid morphology, phylogeny, and biogeography.
Masters Thesis. North Carolina State University. 180 pp.
Eddy and Clarke, 2008. A re-evaluation of a well-preserved skull of Acrocanthosaurus
atokensis supports its carcharodontosaurid affinities. Journal of Vertebrate
Paleontology. 28(3), 73A-74A.
Franzosa and Rowe, 2008 online. Acrocanthosaurus atokensis, Digital Morphology. http://digimorph.org/specimens/Acrocanthosaurus_atokensis/
Bates, 2009. Predicting speed, gait and metabolic cost of locomotion in the
large predatory dinosaur Acrocanthosaurus using evolutionary robotics.
Journal of Vertebrate Paleontology. 29(3), 59A.
Eddy and Clarke, 2011. New information on the cranial anatomy of Acrocanthosaurus
atokensis and its implications for the phylogeny of Allosauroidea (Dinosauria:
Theropoda). PLoS ONE. 6(3), e17932.
Main, Noto and Scotese, 2011. New theropod material from the Cretaceous (Cenomanian)
Woodbine Formation of North Central Texas: Paleobiogeographic and paleoecological
implications. Journal of Vertebrate Paleontology. Program and Abstracts 2011,
150.
Melstrom and D'Emic, 2011. Acrocanthosaurus atokensis (Dinosauria: Theropoda)
from the Cloverly Formation of Wyoming: Implications for Early Cretaceous North
American ecosystems. Journal of Vertebrate Paleontology. Program and Abstracts
2012, 157.
Bennett, Main, Noto, Anderson and Vranken, 2012. Microvertebrate paleoecology,
wildfires and biodiversity of coastal Appalachia in the Cretaceous (Cenomanian)
Woodbine Formation at the Arlington archosaur site, North Texas. Journal of
Vertebrate Paleontology. Program and Abstracts 2012, 63.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
D'Emic, Melstrom and Eddy, 2012. Paleobiology and geographic range of the large-bodied Cretaceous theropod dinosaur Acrocanthosaurus atokensis. Palaeogeography, Palaeoclimatology, Palaeoecology. 333-334, 13-23.
Bourke, Witmer, Ridgely, Porter and Zanno, 2016. Airflow simulations in the antorbital sinus of Acrocanthosaurus.
Testing the potential for theropod paranasal sinuses to function as
accessory cooling structures. Journal of Vertebrate Paleontology.
Program and Abstracts, 101-102.
A? sp. indet. (Burge, 1996)
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Material- teeth, bone fragments
Reference- Burge, 1996. New dinosaur discoveries in the Lower Cretaceous
of Southeastern Utah. Proceedings of Southwest Paleontological Society and Mesa
Southwest Museum, Mesa, Arizona. 4, 85-105.
A? sp. indet. (Cranwell, 2003)
Late Early Cretaceous
Shellenberger Canyon Formation, Arizona, US
Material- caudal centra
Reference- Cranwell, 2003. New evidence of dinosaurs from the Shellenberger
Canyon Formation (Lower Cretaceous) of southeastern Arizona, USA. Southwest
Paleontological Symposium 2002, Guide to Presentations, Mesa Southwest Museum,
unnumbered.
Veterupristisaurus Rauhut,
2011
V. milneri Rauhut, 2011
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Holotype- (MB R 1938; = ST 270) (~10 m) mid caudal vertebra (123 mm)
Paratype-....(MB R 2166; = ST 757) two fused mid caudal vertebrae (91,
~90 mm) fused with chevron fragment
Diagnosis- (after Rauhut, 2011) mid caudal spinoprezygapophyseal lamina
extends anteriorly to midwidth of the base of the prezygapophysis (rather than
its lateral margin); mid caudal spinoprezygapophyseal lamina flanked laterally
by short, parallel lamina extending from lateral margin of prezygapophysis posteriorly.
Comments- These vertebrae were originally referred to Ceratosaurus
roechlingi by Janensch (1925), though Rauhut (2011) described them as a
new taxon of carcharodontosaurid most closely related to Acrocanthosaurus
based on the shared strong anterolateral ridge on the mid caudal transverse
processes.
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. 1(supp. 7), 1-99.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
unnamed possible carcharodontosaurid (Young, 1948)
Barremian-Aptian?, Early Cretaceous
Jiaguan Formation?, Songkanzhen, Guizhou, China
Material- (IVPP V361) (~18 m)
axis (150 mm), about ten rib fragments, two incomplete fused sacral
vertebrae, proximal (?)ischium, fragments
Comments- Discovered in 1944
"about 300 meters NN.W. from the military post at Chingkangsao near a
local town called Sungkan, 43 km from the northern gate of Tungtze
Hsien right along the Chungking-Kweiyang High Way." Sungkan town
is now Songkanzhen, Tungtze Hsien is now Tongzi County, and the
Chongqing-Guiyang Highway is China National Highway 210 (G210).
Young stated it was "found from the basal part of the Cretaceous Beds,
lying about 150 meters above the contact with the Jurassic Beds."
While no other Cretaceous fossils have been reported in Tongzi County
(besides "a corprolite of fish" associated with IVPP V361), about 54 km
north is the Lotus tracksite placed in the Jiaguan Formation, which is
also where tracksites 95 km further southwest in Guizhou like Baoyuan
are assigned. This may be the most appropriate assignment for the
Songkanzhen locality as both it and Baoyuan are equally separated from
the main Sichuan Basin.
Young assigned IVPP V361 to Sauropoda gen. et sp. indet. based on its
large size, "the shape of the centrum, the massiveness of the neural
arch and the dorsal spine", considering it a posteriormost dorsal based
on "the weak opisthocoelus character of the centrum [untrue, actually
amphicoelous] and the anterior position of the pleuro-central cavity,
the shortness of the diapophyses as well as the poor development or
disappearence of some laminae." Noting the neural spine was quite
unlike any known sauropod including Chinese forms (Omeisaurus, Tienshanosaurus, Euhelopus), Young proposed the most similarity to "posterior dorsal vertebrae of Apatosaurus louisae
[which] show a rather flat appearence in the anterior end of the
centrum, low position of the diapophyses and even the dorsal spine of
the dorsal vertebra no. nine shows broader and thicker tip, (but not
anticlinal), but the prominent anterior and posterior as well as the
lateral ridges preventing any further companson." He concluded
"very probably that we have to do with a new peculiar Sauropod."
More recently, Averianov and Sues (2017) proposed "The simple,
paddle-shaped neural spine of the anterior caudal [in Qingxiusaurus]
resembles that of an isolated sauropod vertebra from the Cretaceous of
the neighboring Guizhou Province (Young, 1948), which may belong to the
same or a closely related taxon." Identification of the isolated
element in Qingxiusaurus as a
caudal neural spine notwithstanding, it strongly differs from IVPP V361
in narrowing transversely from near its apex to a very slender base,
being dorsoventrally convex anteriorly in lateral view, having a
transversely flat posterior surface, and having a postspinal lamina at
its base.
Mortimer (online 2015) noted a resemblence to the axis of Acrocanthosaurus in the "spoon-like neural spine ... [and] lack of neural spine laminae", which will be elaborated upon here. Unlike the Acrocanthosaurus
specimen (SMU 74646 3B-1), the axial intercentrum and odontoid are not
fused to the axis in IVPP V361, which explains the absent parapophyses
are these are on the intercentrum in Acrocanthosaurus.
The centra are similar in shape with pleurocoels in the same position
(note the absence of illustrated foramina in Young's specimen may only
indicate preparation was incomplete as the neural canal is drawn as if
filled as well) and a median ventral keel. The "protuberance
directing posteriorly and overhanging the anterior part of the
pleuro-central cavity, a feature also not observed in other Sauropods"
corresponds to the diapophysis in Acrocanthosaurus.
Young's "A. L., accessory lamina (probably infraprezygapophysial
lamina)" that runs ventrolaterally in anterior view is like the
intraprezygapophyseal lamina in Acrocanthosaurus.
The supposed diapophyses are postzygapophyses mostly composed of their
epipophyses, much of which has been broken off. The neural spines
are both very tall and saddle-shaped (anterior edge concave in lateral
view, anterior surface transversely convex, posterior surface
transversely concave) with no laminae, an anterior median ridge and
thickened edges apically. Differences from Acrocanthosaurus
include a shallower ventral median concavity on the centrum in lateral
view, a taller neural arch between the centrum and neural spine,
smaller postzygapophyses angled ventrally, a more vertical neural spine
and a larger projection (of the interspinous ligament?) located at the
anterior base of the neural spine. Other differences are less
clear due to the damage and distortion of IVPP V361, such as the
supposed fossa between what would be the centropostzygapophyseal and
postzygodiapophyseal laminae, and the basally compressed postspinal
fossa. Perhaps the greatest problem with IVPP V361 being a
theropod axis is its size, being 83% longer than SMU 74646 3B-1 (whose
axis minus the intercentrum and odontoid is about 82 mm) with a centrum
85% wider posteriorly. This leads to a total length estimate of
18 meters based on scaling SMU 74646 femoral length to more complete Acrocanthosaurus skeleton NCSM 14345.
Of the two other elements described, the sacral characters are not
sufficient to distinguish theropod from sauropod. The
appendicular fragment is figured in two views and interpreted as a
distal sauropod pubis. If it is a theropod element instead, a
pelvic identification still seems most likely. One possibility is
a proximal ischium with the shorter articular surface being a pubic
peduncle, the grooved surface being acetabular as occurs in some
theropods, and the small area at the top of the figure being the ilial
peduncle. If so, Acrocanthosaurus
differs in having a distinct ilial peduncle, a convex posterior edge to
the ilial peduncle, and more concave edges to the pubic peduncle and
acetabulum.
References- Young, 1948. Notes on the occurrence of sauropod remains from N. Kweichow, China. Science Record. 2(2), 200-206.
Mortimer, online 2015. https://equatorialminnesota.blogspot.com/2015/11/the-kweichow-sauropod.html?showComment=1449558212299#c7366124702472127894
Averianov and Sues, 2017 (online 2016). Review of Cretaceous sauropod
dinosaurs from central Asia. Cretaceous Research. 69, 184-197.
unnamed clade (Carcharodontosaurus saharicus <- Acrocanthosaurus
atokensis)
Comments- This subset of carcharodontosaurids is characterized by wrinkled
tooth enamel, among other characters.
undescribed Carcharodontosauridae (Torices, Barroso-Barcenilla, Cambra-Moo,
Perez-Garcia and Segura, 2010)
Mid-Late Cenomanian, Late Cretaceous
Upper Arenas de Utrillas Formation, Spain
Material- teeth
Comments- These are reported to "have rough enamel and arcuate wrinkles."
Reference- Torices, Barroso-Barcenilla, Cambra-Moo, Perez-Garcia and
Segura, 2010. The new Cenomanian vertebrate site Algora; (Guadalajara, Spain).
Journal of Vertebrate Paleontology. Program and Abstracts 2010, 176A.
undescribed carcharodontosaurid (Lu, Xu, Jiang, Jia, Li, Yuan, Zhang
and Ji, 2009)
Aptian-Albian, Early Cretaceous
Haoling Formation, Henan, China
Material- (41HIII-0093) incomplete lateral tooth
(41HIII-0094) incomplete lateral tooth
(41HIII-0095) anterior tooth (65 mm)
(41HIII-0096) incomplete lateral tooth
Comments- While originally assigned to the Mangchuan Formation, Xu et
al. 2012 split this into three new formations and revised their ages.
Reference- Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009. A preliminary
report on the new dinosaurian fauna from the Cretaceous of the Ruyang Basin,
Henan province of Central China. Journal of the Paleontological Society of Korea.
25(1), 43-56.
unnamed Carcharodontosauridae (Rauhut, 1999)
Cenomanian, Late Cretaceous
Wadi Milk Formation, Sudan
Material- (Vb-607) proximal caudal centrum (137 mm)
(Vb-717) proximal caudal centrum (136 mm)
?(Vb-718) pedal phalanx III-2 (105 mm)
?(Vb-849) distal tarsal III, proximal metatarsal III
(Vb-870) mid caudal centrum (130 mm)
(Vb-871) mid caudal vertebra (136 mm)
Comments- Pleurocoels in Vb-607 and Vb-871 may indicate referral to Carcharodontosaurus.
Vb-871 is from a different taxon than Vb-607, Vb-717 and Vb-870, based on the
deeper ventral groove, less prominent ventral keel, and more distally developed
pleurocoels. The limb elements are only tentatively referred to Carcharodontosauridae,
based on similarity to allosauroids.
References- Werner, 1991. Aspects on terrestrial Upper Cretaceous ecosystems
of Egypt and Northern Sudan. Fifth Symposium on Mesozoic Terrestrial Ecosystems
and Biota, extended abstracts. Contributions from the Paleontological Museum,
University of Oslo. 364, 71-72.
Werner, 1993. Late Cretaceous continental vertebrate fauns of Niger and Northern
Sudan. In Thorweihe and Schandelmier (eds.). Geosicentific Research in Northeast
Africa. Proceedings of the International Conference on Geoscientific Research
in Northeast Africa. 401-405.
Werner, 1994. Die kontinentale Wirbeltierfauna aus der unteren Oberkreide des
Sudan (Wadi Milk Formation) [The continental vertebrate fauna of the lower Upper
Cretaceous of Sudan (Wadi Milk Formation)]. In Kohring and Martin (eds.). Miscellanea
Palaeontologica 3: Festschrift Bernard Krebs. Berliner Geowissenschaften Abhandlungen,
Reihe E. 13, 221-249.
Rauhut, 1999. A dinosaur fauna from the Late Cretaceous (Cenomanian) of Northern
Sudan. Palaeontologia Africana. 35, 61-84.
unnamed Carcharodontosauridae (Vickers-Rich, Rich, Lanus, Rich and Vacca,
1999)
Aptian, Early Cretaceous
Cerro Castano Member of the Cerro Barcino Formation, Chubut, Argentina
Material- (MPEF-PV 1160-1167, 1169) nine tooth fragments
(MPEF-PV 1168) dentary tooth (69.6 mm)
Comments- May be referrable to Tyrannotitan, based on provenance.
References- Vickers-Rich, Rich, Lanus, Rich and Vacca, 1999. 'Big Tooth'
from the Early Cretaceous of Chubut Province, Patagonia: A possible carcharodontosaurid.
In Tomida, Rich, and Vickers-Rich (eds.). Proceedings of the Second Gondwanan
Dinosaur Symposium. National Science Museum Monographs. 15, 85-88.
Rich, Vickers-Rich, Novas, Cuneo, Puerta and Vacca, 2000. Theropods from the
"Middle" Cretaceous Chubut Group of the San Jorge sedimentary basin,
Central Patagonia: A preliminary note. GAIA. 15, 111-115.
undescribed Carcharodontosauridae (Rich, Vickers-Rich, Novas, Cuneo,
Puerta and Vacca, 2000)
Albian-Cenomanian, Early Cretaceous-Late Cretaceous
Bayo Overo Member of the Cerro Barcino Formation, Chubut, Argentina
Material- (MPEF-PV 1171-1174) tooth, three tooth fragments (Rich et al.,
2000)
(probably at least six individuals) fifty-five teeth (Canale et al., 2014)
Comments- From the same locality as "Megalosaurus" inexpectatus,
so may be referrable to that taxon.
References- Rich, Vickers-Rich, Novas, Cuneo, Puerta and Vacca, 2000.
Theropods from the "Middle" Cretaceous Chubut Group of the San Jorge
sedimentary basin, Central Patagonia: A preliminary note. GAIA. 15, 111-115.
Canale, Carballido, Otero, Canudo and Garrido, 2014. Carcharodontosaurid teeth
associated with titanosaur carcasses from the Early Cretaceous (Albian) of the
Chubut Group, Chubut Province, Patagonia, Argentina. Jornadas Argentinas de
Paleontologia de Vertebrados. Ameghiniana. 51(6) suplemento, 6.
undescribed carcharodontosaurid (Calvo, Rubila and Moreno, 1999)
Early Cenomanian, Late Cretaceous
Candeleros Formation of Rio Limay Subgroup, Neuquen, Argentina
Material- maxilla, dentary, cervical vertebrae, dorsal vertebrae, sacral
vertebrae, caudal vertebrae, chevrons, ilia, pubis, femur, tibia, pedal elements
Reference- Calvo, Rubila and Moreno, 1999. Report of a new theropod dinosaur
from northwestern Patagonia. XV Jornadas Argentinas de Paleontolog�a
de Vertebrados. Ameghiniana. 36(4), 7R.
unnamed possible carcharodontosaurid (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
Mid Cenomanian-Turonian, Late Cretaceous
Lower Bajo Barreal Formation, Chubut, Argentina
Material- (UNPSJB-PV 969) tooth (34.2x15.2x7.4 mm)
Comments- Casal et al. (2009)
described this as a carcharodontosaurid tooth, but Lamanna et al.
(2020) stated "note, however, that this tooth may instead pertain to an
abelisauroid [LMI pers. obs.])."
References- Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009. Theropod teeth
(Dinosauria: Saurischia) from the Bajo Barreal Formation, Upper
Cretaceous, Chubut Province, Argentina. Geobios. 42, 553-560.
Lamanna, Casal, Martinez and Ibiricu, 2020. Megaraptorid (Theropoda:
Tetanurae) partial skeletons from the Upper Cretaceous Bajo Barreal
Formation of central Patagonia, argentina: Implications for the
evolution of large body size in Gondwanan megaraptorans. Annals of
Carnegie Museum. 86(3), 255-294.
undescribed Carcharodontosauridae (Canudo, Salgado, Barco, Bolatti and
Ruiz-Omenaca, 2004)
Late Cenomanian-Early Turonian, Late Cretaceous
Cerro Lisandro Formation, Rio Negro, Argentina
Material- (Endemas PV-2) tooth
teeth
Reference- Canudo, Salgado, Barco, Bolatti and Ruiz-Omenaca, 2004. Dientes
de dinosaurios teropodos y sauropodos de la Formacion Cerro Lisandro (Cenomaniense
superior-Turoniense inferior, Cretacico superior) en Rio Negro (Argentina).
Geo-Temas. 6(5), 31-34.
undescribed Carcharodontosauridae (Novas, Martinez, de Valais and Ambrosio,
1999)
Turonian, Late Cretaceous
Mata Amarilla Formation, Santa Cruz, Argentina
Material- teeth
Reference- Novas, Martinez, de Valais and Ambrosio, 1999. Nuevos registros
de Carcharodontosauridae (Dinosauria, Theropoda) en el Cret�cico de Patagonia.
Ameghiniana. 36, 17R.
unnamed carcharodontosaurid (Veralli and Calvo, 2003)
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of Rio Neuquen Subgroup, Neuquen, Argentina
Material- (MUCPv 381, 384, 386, 387, 391) ten teeth
References- Veralli and Calvo, 2003. New findings of carcharodontosauid
teeth on Futalognko quarry (Upper Turonian), north Barreales Lake, Neuqu�n,
Argentina. Ameghiniana. 40(4), 74R.
Veralli and Calvo, 2004. Dientes de ter�podos carcharodontos�uridos
del Turoniano superior-Coniaciano inferior del Neuqu�n, Patagonia, Argentina.
Ameghiniana. 41(4), 587-590.
undescribed carcharodontosaurid (Martinelli and Forasiepi, 2004)
Campanian-Maastrichtian, Late Cretaceous
Allen Formation, Rio Negro, Argentina
Material- (MACN-PV RN 1086) tooth
Reference- Martinelli and Forasiepi, 2004. Late Cretaceous vertebrates
from Bajo de Santa Rosa (Allen Formation), R�o Negro province, Argentina,
with the description of a new sauropod dinosaur (Titanosauridae). Revista del
Museo Argentino de Ciencias Naturales. 6(2), 257-305.
undescribed carcharodontosaurid (Medeiros and Schultz, 2002)
Cenomanian, Late Cretaceous
Alcantara Formation of the Itapecuru Group, Brazil
Material- teeth
Comments- Referred to Carcharodontosaurus sp. by Medeiros and
Schultz (2002), but this is unlikely given the location.
References- Medeiros and Schultz, 2002. The dinosaurian fauna of "Laje
do Coringa", Middle Cretaceous of northeastern Brazil. Arquivos do Museu
Nacional, Rio de Janeiro. 60(3), 155-162.
Castro, Bertini, Santucci and Medeiros, 2005. Fossils from the Coroata Locality,
undifferentiated geological unity, Itapecuru Group, Lower/Middle Albian from
the Sao Luis-Grajau Basin, Maranhao State, North/Northeastern Brazil. In Kellner,
Henriques and Rodriguesn (eds.). II Congresso Latino-Americano de Paleontologia
de Vertebrados, Rio De Janeiro Museu Nacional. Boletim de Resumos. 75-76.
Elais, Bertini and Medeiros, 2005. Review of the occurrences concerning isolated
amniotes teeth, in the Cretaceous deposits from the Maranhao State. In Kellner,
Henriques and Rodriguesn (eds.). II Congresso Latino-Americano de Paleontologia
de Vertebrados, Rio De Janeiro Museu Nacional. Boletim de Resumos. 99-100.
undescribed Carcharodontosauridae (Kellner and Campos, 1998)
Turonian-Santonian, Late Cretaceous
Adamantina Formation of the Bauru Group, Brazil
Material- (MMR/UFU-PV 005) tooth (Candeiro et al., 2006)
(UFRJ-DG 379-Rd) tooth (Candeiro et al., 2004)
References- Kellner and Campos, 1998. Review of Cretaceous theropods
and sauropods from Brazil. Journal of Vertebrate Paleontology. 18(3), 55A.
Silva and Kellner, 1999. Novos dentes de Theropoda do Cretaceo continental do
Brasil. Paleontologia em Destaque, Boletim Informativo da Sociedade Brasileira
de Paleontologia. 14(26).
Candeiro, Abranches, Abrantes, Avilla, Martins, Moreira, Torres and Bergqvist,
2004. Dinosaurs remains from western S�o Paulo state, Brazil (Bauru Basin,
Adamantina Formation, Upper Cretaceous). Journal of South American Earth Sciences.
18, 1-10.
Candeiro, Martinelli, Avilla and Rich, 2006. Tetrapods from the Upper Cretaceous
(Turonian-Maastrichtian) Bauru Group of Brazil: A reappraisal. Cretaceous Research.
27, 923-946.
Candeiro, Santos, Rich, Marinho and Oliveira, 2006. Vertebrate fossils from
the Adamantina Formation (Late Cretaceous), Prata paleontological district,
Minas Gerais State, Brazil. Geobios. 39, 319-327.
undescribed Carcharodontosauridae (Candeiro, Bergqvist, Novas and Currie,
2004)
Late Maastrichtian, Late Cretaceous
Serra da Galga Formation of the Bauru Group, Brazil
Material- (CPP 124, 127, 129a, 152, 156, 197, 199, 200, 208, 216, 241,
375/1, 376, 447-449, 474, 475) eighteen teeth
Comments- In 2021 the Serra da Galga and Ponte Alta Members of the Marilia Formation were recognized as the Serra da Galga Formation.
References- Candeiro, Bergqvist, Novas and Currie, 2004. Theropod teeth
from the Marilia Formation (Upper Maastrichtian), Minas Gerais state, Brazil.
Journal of Vertebrate Paleontology. 24(3), 204A.
Candeiro, Martinelli, Avilla and Rich, 2006. Tetrapods from the Upper Cretaceous
(Turonian-Maastrichtian) Bauru Group of Brazil: A reappraisal. Cretaceous Research.
27, 923-946.
Candeiro, Currie and Bergqvist, 2012. Theropod teeth from the Mar�lia
Formation (Late Maastrichtian) at the paleontological site of Peir�polis
in Minas Gerais State, Brazil. Revista Brasileira de Geoci�ncias. 42(2),
323-330.
unnamed clade (Labocania anomala + Shaochilong maortuensis)
Diagnosis- very thick frontal; supratemporal fossa restricted from frontal;
sagittal crest on frontal; lack of helical groove on quadrate condyle.
Comments- Labocania and Shaochilong appear to be each others
closest relatives, a possibility originally noted by Molnar (1974), and later
elaborated by Chure (2000). The clade remains unnamed however. The taxa fall
outside Tyrannoraptora in a modified version of Senter's (2007) matrix, as Shaochilong
lacks a palatal shelf, nasal fusion or transversely convex nasals, and has fused
interdental plates. While the triangular obturator process of Labocania
and the subcondylar recesses of Shaochilong are similar to coelurosaurs,
these are also present in some carcharodontosaurids.
Labocania Molnar, 1974
L. anomala Molnar, 1974
Late Campanian, Late Cretaceous
El Gallo Formation, Mexico
Diagnosis- pneumatic quadrate; oblique ridge on posterior face of quadrate;
lateral dentary shelf.
Holotype- (IGM 5307; = LACM 20877) (~7.5 m; ~1.2 tons) (skull ~660 mm) partial maxilla,
distal quadrate, frontal, dentary fragment, premaxillary teeth, maxillary teeth,
chevron, ilial fragment, proximal ischium (~750 mm), shaft of pubis, nearly complete metatarsal
II (<507 mm), phalanx III-2 (~115 mm), long bone fragments
Comments- Discovered in 1970, this taxon was originally compared to Indosaurus,
Shaochilong and an unnamed partial dentary from the Kelaza Formation
described by Young and Sun (1957) (IVPP V903). Similarities to tyrannosaurids
were also noted. Paul (1988) referred it to his Allosauridae, which was paraphyletic
to tyrannosaurids. Lamanna and Smith reexamined this species in 1998 and found
the specimen to be more fragmentary than expected. A paper may be published
soon. They conclude it is a tyrannosaurid, although it may not be valid. Chure
(2000) referred it to Holtz's tyrannosaur + ornithomimosaur + troodontid clade,
and thought it was most closely related to Shaochilong. Another so far
unpublished possibility is a relationship to carcharodontosaurids, as Mapusaurus
and Giganotosaurus have a pneumatic quadrate, lateral dentary shelf,
proximolateral ischial fossa and triangular obturator process. This would coincide
with Brusatte et al.'s (2009) recent assignment of Shaochilong to that
clade.
References- Young and Sun, 1957. Note on a fragmentary carnosaurian mandible
from Turfan, Sinkiang. Vertebrata PalAsiatica. 1(2), 2027-2036.
Molnar, 1974. A distinctive theropod dinosaur from the Upper Cretaceous of Baja
California (Mexico). Journal of Paleontology. 48(5), 1009-1017.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Brusatte, Benson, Chure, Xu, Sullivan and Hone, 2009. The first definitive carcharodontosaurid
(Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids.
Naturwissenschaften. 96(9), 1051-1058.
Rivera-Sylva and Carpenter, 2014. Mexican saurischian dinosaurs. In
Rivera-Sylva, Carpenter and Frey (eds.). Dinosaurs and Other Reptiles
from the Mesozoic of Mexico. Indiana University Press. 143-155.
Ram�rez-Velasco and Hern�ndez-Rivera, 2015. Diversity of Late
Cretaceous dinosaurs from Mexico. Bolet�n Geol�gico y Minero. 126(1),
63-108.
Shaochilong Brusatte,
Benson, Chure, Xu, Sullivan and Hone, 2009
= “Alashansaurus" Chure, 2000
S. maortuensis (Hu, 1964) Brusatte, Benson, Chure, Xu, Sullivan
and Hone, 2009
= Chilantaisaurus maortuensis Hu, 1964
= "Alashansaurus" maortuensis (Hu, 1964) Chure, 2000
Turonian, Late Cretaceous
Maortu, Lower Ulanhsuhi Formation of Dashigou Group, Inner Mongolia, China
Lectotype- (IVPP V2885.1) (~5-6 m; ~500 kg; adult) (skull ~580 mm) incomplete
braincase
....(IVPP V2885.2) partial nasal, frontals (93.3 mm), parietals (22 mm)
Paralectotypes- ....(IVPP V2885.3) quadrates (143 mm)
....(IVPP V2885.4) maxillae (one incomplete, one fragmentary and lost; ~350
mm)
....(IVPP V2885.5) incomplete axis (~70 mm)
?...(IVPP V2885.6) incomplete proximal caudal vertebra (72 mm), two proximal
caudal vertebrae (lost)
?...(IVPP V2885.7) mid caudal vertebra (85 mm), distal caudal centrum (85 mm),
distal caudal centrum (90 mm)
Diagnosis- (after Hu, 1964) twelve maxillary teeth.
(after Chure, 1998) paradental groove on medial surface of maxilla absent; large
cylindrical pneumatic cavity in posterior nasals; deep sagittal crest on frontals
and parietals.
(after Chure, 2000) no maxillary palatal shelf.
(after Brusatte et al., 2009) maxillary antorbital fossa reduced in extent and
nearly absent; deep, dorsoventrally oriented grooves located dorsally on maxillary
interdental plates; large pneumatic foramen at anterodorsal corner of dorsal
tympanic recess of prootic.
Other diagnoses- Hu (1964) also diagnosed maortuensis by its large
occipital condyle (compared to the foramen magnum), which is similar to other
carcharodontosaurids. His last two characters- small skull and small quadrate,
are too vague to evaluate.
Of the supposedly diagnostic characters listed by Chure (1998), the fused maxillary
interdental plates, anteriorly limited supratemporal fossae, declined paroccipital
processes and highly pneumatized and shortened basicranium are typical of carcharodontosaurids.
Contra Chure, a paraquadrate foramen was present, and the maxillary interdental
plates are not small. The frontal sagittal crest is shared with Labocania.
Chure (2000) listed additional diagnostic characters, including ones that are
typical of carcharodontosaurids- thick, flat frontals; subnarial process of
frontals forms a deep trough; short axis. The absent distal quadrate groove
is shared with Labocania, while the frontals aren't unfused and the axial
neural spine is posterodorsally inclined.
Comments- The material was discovered in 1960. Hu (1964) placed this and Chilantaisaurus tashuikensis
in the same genus based on dental and caudal similarities. The caudals of Chilantaisaurus
are only doubtfully referred and belong to the proximal part of the series,
while those of Shaochilong are more distal. Also, the tooth of Chilantaisaurus
is similarly doubtfully referred and comparison with the exposed tooth of Shaochilong
is not useful. Because of this, Chure (1998) separated the two species, and
later (2000) created a new genus for maortuensis in his unpublished thesis-
"Alashansaurus" (incorrectly spelled "Alshansaurus" by Brusatte
et al., 2010). The name was published by Glut (2003), but the latter reference
includes a caveat to prevent it from being an official taxonomic source, leaving
the possibility of Chure being its official describer once his thesis' contents
are published. This happened in 2009, when Chure coauthored Brusatte et al.
(2009), naming the genus Shaochilong instead.
After Hu's assignment to the Megalosauridae, Molnar (1974) noted frontal and
quadrate similarities to Labocania. Both Paul (1988) and Molnar et al.
(1990) assigned it to a paraphyletic Allosauridae, closer to tyrannosaurids
than Allosaurus. Chure (1998) noted it shared characters with Labocania,
tyrannosaurs, troodontids and dromaeosaurids. He later (2000) assigned it to
Holtz's tyrannosaur + ornithomimosaur + troodontid clade based on the proximolateral
ischial scar in the related Labocania, and to the Tyrannosauroidea based
on the highly pneumatized basicranium and short and deep braincase. He finds
it is most closely related to Labocania, which I concur with based on
characters described above. Brusatte et al. (2009) entered it into Smith et
al.'s (2007) matrix, finding it to be a carcharodontosaurid outside of Carcharodontosaurinae
and related to Tyrannotitan.
References- Hu, 1964. Carnosaurian remains from Alashan, Inner Mongolia.
Vertebrata PalAsiatica. 8, 42-63.
Molnar, 1974. A distinctive theropod dinosaur from the Upper Cretaceous of Baja
California (Mexico). Journal of Paleontology. 48(5), 1009-1017.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 169-209.
Chure, 1998. "Chilantaisaurus" maortuensis, a large maniraptoran
theropod from the Early Cretaceous (Albian) of Nei Mongol, PRC. Journal of Vertebrate
Paleontology. 18(3), 33A-34A.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Glut, 2003. Dinosaurs - The Encyclopedia - Supplement 3. McFarland Press, Jefferson,
NC. 726 pp.
Brusatte, Benson, Chure, Xu, Sullivan and Hone, 2009. The first definitive carcharodontosaurid
(Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids.
Naturwissenschaften. 96(9), 1051-1058.
Brusatte, Chure, Benson and Xu, 2010. The osteology of Shaochilong maortuensis,
a carcharodontosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Asia.
Zootaxa. 2334, 1-46.
Carcharodontosaurinae Stromer, 1931
vide Brusatte and Sereno, 2008
Definition- (Carcharodontosaurus saharicus + Giganotosaurus
carolinii) (Brusatte and Sereno, 2008)
"Megalosaurus" chubutensis
Corro, 1974
Cenomanian-Turonian, Late Cretaceous
Cerro Castillo Formation, Chubut, Argentina
Holotype- (MACN 18.189) tooth (~85 mm)
Comments- Poblete and Calvo (2004) assign chubutensis to Carcharodontosauridae,
based on marginal wrinkles, serration density and a distal carina which descends
in a zigzag pattern in lingual view. They consider it indeterminate. Carrano
et al. (2012) stated it resembled abelisaurids, but did not list shared characters.
Reference- Corro, 1974. Un nuevo megalosaurio (Carnosaurio) del Cretacico
de Chubut (Argentina). Comunicaci�n del Museo Argentino de Ciencias Naturales
Bernardino Rivadavia. 1, 37-44.
Poblete and Calvo, 2004. "Megalosaurus chubutensis" del Corro:
un posible Carcharodontosauridae del Chubut. Ameghiniana. 41(4), 59R-60R.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
"Megalosaurus" ingens
Janensch, 1920
= Ceratosaurus ingens (Janensch, 1920) Paul, 1988
Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
Lectotype- (MB R 1050) tooth (120 mm)
Paralectotypes- (MB R 1060) tooth
(MB R 1069) tooth
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Paralectotypes- (MB R 1054) tooth
(MB R 1057) tooth
(MB R 1058) tooth
(MB R 1061) tooth
(MB R 1067) tooth
Callovian-Oxfordian, Middle Jurassic-Late Jurassic
Lower Dinosaur Member of the Tendaguru Formation, Tanzania
Paralectotypes- (MB R 1053) tooth
(MB R 1064) tooth
Callovian-Tithonian, Middle Jurassic-Late Jurassic
Tendaguru Formation, Tanzania
Paralectotypes- (MB R 1082) tooth
?(MB R coll.) thirteen teeth, tooth fragments
Comments- Though Janensch (1925) referred 13 more teeth and tooth fragments
to the taxon, these are excluded by Rauhut (2011) as not sharing the exact same
combination of characters. Rauhut did note there are Tendaguru teeth that are
similar except for the absence of enamel wrinkles, which are individually variable
in many other taxa. While this species has been referred to both Megalosaurus
(Janensch, 1920) and Ceratosaurus (Paul, 1988), Rauhut (2011) noted it
differs from these taxa. Specifically, Megalosaurus differs in having
a more strongly recurved crown tip, finer serrations, mesial serrations that
ends well above the crown-root junction, and lacks down-pointing grooves at
the bases of the serrations. Ceratosaurus also lacks the latter grooves,
has strongly transversely flattened crowns, pronounced flat or even slightly
concave areas adjacent to the carinae, and never shows enamel wrinkles. Rauhut
(1995) noted similarity with Carcharodontosaurus in the interdenticle
grooves, and suggested it may be a carcharodontosaurid. He later (2011) made
the same suggestion, this time based on the combination of only slightly recurved
crowns, basally directed grooves at the base of the serrations, mesial carina
extends to the base of the crown, and marginal, apically curved enamel wrinkles
along the carinae. If so, these characters are only shared with carcharodontosaurines
within that family. Note Madsen and Welles (2000) incorrectly attributed the
combination Ceratosaurus ingens to Rowe and Gauthier (1990) and wrongly
considered it a lapsus calumni, apparently unaware of Paul's book.
References- Janensch, 1920. Ueber Elaphrosaurus bambergi und die
Megalosaurier aus den Tendaguru Schichten Deutsch-Ostafrikas. Sitzungsberichte
der Gesellschaft Naturforschender Freunde zu Berlin. 1920, 225-235.
Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten Deutsch-Ostafrikas.
Palaeontographica. 1(supp. 7), 1-99.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Rowe and Gauthier, 1990. Ceratosauria. In Weishampel, Osm�lska and Dodson
(eds.). The Dinosauria. University of California Press. 151-168.
Rauhut, 1995. Zur systematischen Stellung der afrikanischen Theropoden Carcharodontosaurus
Stromer, 1931, und Bahariasaurus Stromer, 1934. Berliner geowissenschaftliche
Abhandlungen. 16(1), 357-375.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey. 80 pp.
Rauhut, 2006. Theropod dinosaurs from the Late Jurassic of Tanzania and the
origin of Cretaceous Gondwanan theropod faunas. Journal of Vertebrate Paleontology.
26(3), 113A.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
unnamed carcharodontosaurine (Simionescu, 1913)
Late Valanginian, Early Cretaceous
Alimanu Member of the Cernavoda Formation, Romania
Material- (UAIC (SCM1) 615; = Harsova Museum 200) lateral tooth (85.5x29x16.3
mm)
Comments- This tooth was described by Simionescu (1913) and compared
to Erectopus (then Megalosaurus superbus). Huene (1926) doubted
it was the same species due to age differences, but did continue to call it
Erectopus aff. superbus. Csiki-Sava et al. (2016) redescribed
the specimen in detail and determined it is strongly supported as a carcharodontosaurine
and resembles Carcharodontosaurus more than giganotosaurines in two characters-
roughly straight distal margin of crown; pronounced marginal undulations in
enamel that are easily visible in normal light.
References- Simionescu, 1913. Megalosaurus aus der Unterkreide
der Dobrogea. Centralblatt f�r Mineralogie, Geologie, und Palaeontologie.
1913, 686-687.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
Csiki-Sava, Brusatte and Vasile, 2016. "Megalosaurus cf. superbus"
from southeastern Romania: The oldest known Cretaceous carcharodontosaurid (Dinosauria:
Theropoda) and its implications for earliest Cretaceous Europe-Gondwana connections.
Cretaceous Research. 60, 221-238.
unnamed carcharodontosaurine (Azevedo, Simbras, Furtado, Candeiro and
Bergqvist, 2012)
Campanian-Maastrichtian, Late Cretaceous
Presidente Prudente Formation, Brazil
Material- (UFRJ-DG409-R) maxillary fragment
Reference- Azevedo, Simbras, Furtado, Candeiro and Bergqvist, 2012. First
Brazilian carcharodontosaurid and other new theropod dinosaur fossils from the
Campanian-Maastrichtian Presidente Prudente Formation, S�o Paulo State,
Southeastern Brazil. Cretaceous Research. 40, 131-142.
Carcharodontosaurus
Stromer, 1931
Diagnosis- (modified after Brusatte and Sereno, 2007) pronounced grooved
sculpturing of nearly the entire lateral surface of the maxilla.
Other diagnoses- Chiarenza and Cau (2016) noted two characters proposed
by Brusatte and Sereno (2007) (large internal carotid and paracondylar pneumatocoels
(pneumatic recesses); deep funnel-shaped basisphenoid fossa) are also present
in Giganotosaurus.
Comments- Contra Weishampel et al. (2004), who listed
Carcharodontosaurus saharicus as being known from the "Continental
intercalaire" of Muhafazat Gharyan, Libya,
References- Stromer, 1931. Ergebnisse der Forschungsreisen Prof. E. Stromers
in den W�sten �gyptens. II. Wirbeltier-Reste der Bahar�jestufe
(unterstes Cenoman). 10. Ein Skelett-Rest von Carcharodontosaurus nov.
gen. Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche
Abteilung, Neue Folge. 9, 1-23.
Brusatte and Sereno, 2007. A new species of Carcharodontosaurus (Dinosauria:
Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal
of Vertebrate Paleontology. 27(4), 902-916.
Smith, Lamanna, Askar, Bergig, Tshakreen, Abugares and Rasmussen, 2010. A large
abelisauroid theropod dinosaur from the Early Cretaceous of Libya. Journal of
Paleontology. 84(5), 927-934.
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco
and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.
C. saharicus (Deperet and Savornin, 1925) Stromer, 1931
= Megalosaurus saharicus Deperet and Savornin, 1925
= Megalosaurus (Dryptosaurus) saharicus (Deperet and Savornin, 1925)
Deperet and Savornin, 1927
= Megalosaurus africanus Deperet and Savornin, 1925 vide Huene, 1956
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Neotype- (UCRC PV12; = SGM-Din 1) (12.79 m) incomplete skull (missing premaxillae,
squamosals, quadratojugals) (~1.42 m) (Sereno et al., 1996)
Referred- ?(CMN 41817) tooth (54x27x12 mm) (Russell, 1996)
?(CMN 41818) tooth (67x36x22 mm) (Russell, 1996)
?(CMN 41819) tooth (69x34x16 mm) (Russell, 1996)
?(CMN 41859) dentary fragment (Russell, 1996)
?(CMN 41908) tooth (30x24x11 mm) (Russell, 1996)
?(CMN 41910) tooth (23x19x6 mm) (Russell, 1996)
?(CMN 50792) cervical vertebra (148 mm) (Russell, 1996)
?(FSAC-KK 02) dentary fragment (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(FSAC-KK 907) partial tooth (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(M-CH-003) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
?(M-TA-032) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
?(MPDM 16) tooth (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(NMB-1673-R) tooth (72.6x36x17.5 mm) (Richter, Mudroch and Buckley, 2013)
?(NMB-1674-R) tooth (67.2x29.3x8.9 mm) (Richter, Mudroch and Buckley, 2013)
?(UCRC PV163) tooth (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(UCRC PV164) tooth (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
? teeth, ilium (Lavocat, 1954)
? teeth (Sadleir, 1998)
Albian, Early Cretaceous
Continental Intercalaire, Algeria
Holotype- ?(lost; syntypes of Megalosaurus saharicus) two maxillary
or dentary teeth
Referred- ?(MNNHN coll.) twelve teeth (to FABL of 42 mm), few mid caudal
vertebrae (~70-100 mm) (Lapparent, 1960)
Cenomanian, Late Cretaceous
Baharija Formation, Egypt
?(IPHG 1912 VIII 68; destroyed) ilium (Stromer, 1934)
(IPHG 1922 X46; holotype of Carcharodontosaurus; destroyed) partial maxilla, maxillary
teeth, nasals, frontals, parietals, supraoccipital, partial exoccipital-opisthotics,
axis, anterior cervical vertebra (100 mm), cervical vertebra, proximal dorsal
rib, proximal caudal vertebra (145 mm), proximal chevron (150 mm), partial chevron,
manual ungual, incomplete pubes (>1 m), partial ischium, femora (1.26 m),
fibula (880 mm) (Stromer, 1931)
?material (Smith et al., 2001)
Cretaceous
Africa
?(FPDM-V6211) fragmentary skull (Azuma, 2005)
Diagnosis- (after Brusatte and Sereno, 2007) laterally protruding ventral
margin of the maxillary external antorbital fossa; deep and ventrally-facing
fossa between
the inner wall of the maxilla and the anteromedial process; deep and dorsoventrally
protruding lacrimal-frontal suture; invaginated anteromedial corner of the supratemporal
fossa; distinct enamel wrinkles on both mesial and distal margins of mesial
(anterior) and mid maxillary crowns.
Comments- Megalosaurus saharicus is based on two teeth from the
Albian Continental Intercalaire of Algeria (Deperet and Sevornin, 1925). Huene
(1956) listed Megalosaurus africanus, incorrectly attributed to Deperet
and Savornin (1925), but this is probably a mistake. Later, Stromer (1931) described
a partial skeleton found in 1914 from the Early Cenomanian Baharija Formation of Egypt and
referred it to this species, creating the genus Carcharodontosaurus for
it. An incomplete skull from the Cenomanian Kem Kem beds of Morocco closely
resembling Stromer's specimen was found in 1995 and reported by Sereno et al. (1996). Brusatte
and Sereno (2007) noted Giganotosaurus has identical teeth to Carcharodontosaurus
saharicus, thus the original Megalosaurus saharicus specimens are
indeterminate. In order to save Carcharodontosaurus saharicus from being
a nomen dubium, they made the incomplete Kem Kem skull the neotype of the species
(Stromer's material is destroyed). Since no diagnostic differences have been
noted between the teeth of Carcharodontosaurus saharicus and Giganotosaurus,
the referral of isolated teeth to any particular derived carcharodontosaurid
taxon is based solely on locality. Chiarenza and Cau (2016) proposed the neotype
differs from the holotype in having ventrally flattened maxillary interdental
plates, but this seems to be due to damage in the neotype, whose complete third
interdental plate is ventrally pointed.
The ilium (IPHG 1912 VIII 68) referred to Carcharodontosaurus by Stromer
(1934) does not necessarily belong to this taxon, and may be ceratosaurian (?=
Deltadromeus or Bahariasaurus). Twelve teeth and a few mid caudal
vertebrae from the holotype locality were described by Lapparent (1960). The
teeth are described as having enamel wrinkles as in derived carcharodontosaurids
and being very similar to the holotype, though the caudals' referral is uncertain.
They are referred here to C. saharicus based on provenance. Russell (1996)
described a maxillary fragment, cervical vertebra and teeth from the Kem Kem
beds of Morocco, which though closely resembling those in Stromer's specimen,
are only referred to the species saharicus based on provenance. The same
can be said for teeth from that locality mentioned by Lavocat (1954) and Sadlier
(1998), though Lavocat's teeth were not only compared to Carcharodontosaurus,
but also Tendaguru taxa and Tyrannosaurus, so may not be referrable to
carcharodontosaurids at all. The supposed maxillary fragment (CMN 41859) was described twice by
Russell, the second time as a dentary fragment of Abelisauridae gen. et
sp. indet.. Ibrahim (2011) agreed it was
actually carcharodontosaurid based on "the deep proportions and
alveolar
morphology" (Ibrahim et al., 2020), with Ibrahim et al. (2017, 2020)
referring it to the coeval Carcharodontosaurus saharcus. Smith et al. (2001) reported new remains of cf.
Carcharodontosaurus from the Baharija Formation of Egypt. Sereno et al.
(1996) referred the specimen of Spinosaurus B and the material of Sigilmassasaurus
to Carcharodontosaurus saharicus, but Ibrahim et al. (2014) have since
demonstrated they belong to Spinosaurus.
References- Deperet and Savornin, 1925. Sur la decouverte d'une faune
de vertebres albiens a Timintoun (Sahara occidental). Comptes Rendus de l’Academic
de Sciences. 181, 1108-1111.
Dep�ret and Savornin, 1927. La faune de reptiles et de poisons albiens
de Timimoun (Sahara alg�rien). Bulletin de la soci�t� g�ologique
de France. 27, 257-265.
Stromer, 1931. Ergebnisse der Forschungsreisen Prof. E. Stromers in den W�sten
�gyptens. II. Wirbeltier-Reste der Bahar�jestufe (unterstes Cenoman).
10. Ein Skelett-Rest von Carcharodontosaurus nov. gen. Abhandlungen der
Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche
Abteilung, Neue Folge. 9, 1-23.
Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers in den W�sten
�gyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes Cenoman). 13.
Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche
Abteilung, Neue Folge. 22, 1-79.
Lavocat, 1954. Sur les dinosauriens du Continental Intercalaire des Kem-Kem
de la Daoura [On the dinosaurs from the Continental Intercalaire of the Kem
Kem of the Doura]. Comptes Rendus 19th Intenational Geological Congress, 1952.
1, 65-68.
Huene, 1956. Palaeontologie und Phylogenie der Niederen Tetrapoden. VEB Gustav
Fischer Verlang, Jena. 716 pp.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du
Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Rauhut, 1995. Zur systematischen Stellung der afrikanischen Theropoden Carcharodontosaurus
Stromer 1931 und Bahariasaurus Stromer 1934. Berliner Geowissenschaften Abhandlungen. 16, 357-375.
Currie, 1996. Out of Africa: Meat-eating dinosaurs that challenge Tyrannosaurus rex. Science. 272(5264), 971-972.
Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt,
Morocco. Bulletin du Museum national d'Histoire naturelle (4e se'r.) 18, 349-402.
Sereno, Dutheil, Iarochene, Larsson, Lyon, Magwene, Sidor, Varricchio and Wilson,
1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation.
Science. 272(5264), 986-991.
Sadleir, 1998. Theropod teeth from the Cretaceous of Morocco. Journal of Vertebrate
Paleontology. 18(3), 74A.
Larsson, 2001. Endocranial anatomy of Carcharodontosaurus saharicus (Theropoda:
Allosauroidea) and its implications for theropod brain evolution. in Tanke and
Carpenter (eds.). Mesozoic Vertebrate Life: New Research inspired by the Paleontology
of Philip J. Currie. Indiana University Press, Bloomington & Indianapolis,
Indiana. 19-33.
Smith, Lamanna, Lacovara, Dodson, Smith, Poole, Giegengack and Attia, 2001.
A giant sauropod dinosaur from an Upper Cretaceous mangrove deposit in Egypt.
Science. 292, 1704-1706.
Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004. Isolated theropod teeth from
the Cenomanian of Morocco and their palaeobiogeographical significance. Revue
de Paleobiologie, Geneve. 9, 143-149.
Azuma, 2005. The Flying Dinosaurs. Fukui Prefectural Dinosaur Museum. 118 pp.
Brusatte and Sereno, 2007. A new species of Carcharodontosaurus (Dinosauria:
Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal
of Vertebrate Paleontology. 27(4), 902-916.
Ibrahim, 2011. Dinosaurs and other fossil vertebrates from the
Cretaceous of southeastern Morocco. PhD thesis, University College
Dublin. 836 pp.
Richter, Mudroch and Buckley, 2013. Isolated theropod teeth from the Kem Kem
Beds (Early Cenomanian) near Taouz, Morocco. Palaontologische Zeitschrift. 87,
291-309.
Ibrahim, Sereno, Dal Sasso, Maganuco, Fabbri, Martill, Zouhri, Myhrvold and
Iurino, 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science.
345(6204), 1613-1616.
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco
and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.
Ibrahim, Sereno and Zouhri, 2017. Les dinosaures du Maroc - aper�u
historique et travaux r�cents. In Zouhri (ed.). Pal�ontologie des
vert�br�s du Maroc: �tat des connaissances. M�moires de la Societ�
g�ologique de France. 2017, 249-284.
Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020. Geology and paleontology of the Upper
Cretaceous Kem Kem Group of eastern Morocco. ZooKeys. 928, 1-216.
C? iguidensis Brusatte
and Sereno, 2007
Cenomanian, Late Cretaceous
Echkar Formation of the Tegama Group, Niger
Holotype- (MNN IGU2) partial maxilla
Paratypes- ?(MNN IGU3) (adult) braincase
?(MNN IGU4) partial lacrimal
?(MNN IGU5) anterior dentary
?(MNN IGU6) tooth
?(MNN IGU7) tooth
?(MNN IGU8) tooth
?(MNN IGU9) tooth
?(MNN IGU10) tooth
? teeth, vertebral fragments
Diagnosis- (after Brusatte and Sereno, 2007) very reduced antorbital
fossa limited to the proximity of the maxillary fenestra; anteromedial maxillary
process that is broadly arched toward the midline; prominent horizontal crest
on the medial aspect of the main maxillary body; braincase excavated by a deep
invaginated fossa on the anterior aspect of the laterosphenoid ala.
Comments- Sereno et al. (2004) noted carcharodontosaurids were associated
with Rugops in the Echkar Formation, and Brusatte and Sereno (2005) briefly
described the new species in an abstract. It was named and described in detail
by Brusatte and Sereno (2007). They also referred a cervical centrum (MNN IG11)
to the species, but it was found loose from the ground and was too immature
to belong to the same individual as MNN IGU3. It is here referred to Spinosaurus,
as Ibrahim et al. (2014) demonstrated cervicals with that morphology belong
to that genus. Chiarenza and Cau (2016) noted the paratypes were not found close
to the holotype, and that the braincase MNN IGU3 lacks several characters present
in Carcharodontosaurus+Giganotosaurus. Furthermore, the two braincase
characters proposed to diagnose Carcharodontosaurus by Brusatte and Sereno
are also present in Giganotosaurus. Thus MNN IGU3 may be a non-carcharodontosaurine.
References- Sereno, Wilson, and Conrad, 2004. New dinosaurs link southern
landmasses in the mid-Cretaceous. Proceedings of the Royal Society of London
B. 271(1546), 1325-1330.
Brusatte and Sereno, 2005. A new species of Carcharodontosaurus (Dinosauria:
Theropoda) from the Cenomanian of Niger and its implications for allosauroid
phylogeny. Journal of Vertebrate Paleontology. 25(3), 40A.
Brusatte and Sereno, 2007. A new species of Carcharodontosaurus (Dinosauria:
Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal
of Vertebrate Paleontology. 27(4), 902-916.
Ibrahim, Sereno, Dal Sasso, Maganuco, Fabbri, Martill, Zouhri, Myhrvold and
Iurino, 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science.
345(6204), 1613-1616.
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco
and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.
C? sp. indet. (Lapparent, 1953)
Albian-Early Cenomanian, Early Cretaceous-Late Cretaceous
Tegama Group, Niger
Material- (MNNHN Td. 2269) lateral tooth fragment (Lapparent, 1960)
(MNNHN coll.; from In Abangarit) two braincase fragments, 137 teeth (anterior
teeth- 80x33 mm, 77x31 mm, 62x28 mm, 64x27 mm, 54x26 mm; lateral teeth- 125x47
mm, 70x45 mm, 105x40 mm, 90x37 mm, 87x36 mm), proximal caudal vertebra (120
mm), distal caudal vertebra, manual phalanx II-2 (60 mm) (Lapparent, 1960)
Comments- This material was not found associated, so could belong to
multiple individuals of several large theropod taxa (Deltadromeus, Bahariasaurus,
Spinosaurus, etc.). Over a hundred of the teeth are carcharodontosaurid,
however. Carcharodontosaurid material may be referrable to Carcharodontosaurus
sp. nov., based on provenance. A pedal ungual from In Abangarit referred to
C. saharicus by Lapparent matches the Spinosaurus B morphotype
that has been shown to be Spinosaurus by Ibrahim et al. (2014). The carcharodontosaurid
tooth fragment MNNHN Td. 2269 was described by Lapparent as being from the younger
Elrhaz Formation, but Taquet (1976) notes it came from a cliff of Albian deposits.
References- Lapparent, 1953. Gisements de dinosauriens dans le "Continental
intercalaire" d'In Abangharit (Saharia meridional). Compte rendu hebdomadaire
des seances de l’Academie des Sciences Paris. 236, 1905-1906.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du
Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Taquet, 1976. G�ologie et Pal�ontologie du Gisement de Gadoufaoua
(Aptien du Niger) [Geology and Paleontology of the Gadoufaoua Locality (Aptian
of Niger)]. Cahiers de Pal�ontologie, Centre National de la Recherche
Scientifique, Paris. 191 pp.
Ibrahim, Sereno, Dal Sasso, Maganuco, Fabbri, Martill, Zouhri, Myhrvold and
Iurino, 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science.
345(6204), 1613-1616.
C? sp. indet. (Schluter and Schwarzhans, 1978)
Late Aptian-Early Albian, Early Cretaceous
Chenini or Oum Ed Diab Member of the Ain el Guettar Formation, Dahar/Jeffara escarpment, Tunisia
Material- (MGGC 21891) incomplete mid caudal vertebra (Fanti et al.,
2014)
?(MGGCTUN9; Morphotype 2) lateral tooth (41x22x14 mm) (Fanti et al., 2014)
?(MGGCTUN14; Morphotype 2) lateral tooth (50x31x19 mm) (Fanti et al., 2014)
?(MGGCTUN19; Morphotype 2) lateral tooth (38x25x15 mm) (Fanti et al., 2014)
?(MGGCTUN39; Morphotype 2) lateral tooth (38x27x16 mm) (Fanti et al., 2014)
?(MGGCTUN41; Morphotype 2) lateral tooth (33x24x12 mm) (Fanti et al., 2014)
?(MGGCTUN42; Morphotype 2) lateral tooth (36x26x14 mm) (Fanti et al., 2014)
?(MGGCTUN46; Morphotype 2) lateral tooth (35x32x18 mm) (Fanti et al., 2014)
?(MGGCTUN72; Morphotype 2) lateral tooth (29x31x17 mm) (Fanti et al., 2014)
?(MGGCTUN86; Morphotype 2) lateral tooth (55x27x12 mm) (Fanti et al., 2014)
?(MGGCTUN87; Morphotype 2) lateral tooth (54x32x15 mm) (Fanti et al., 2014)
?(MGGCTUN111; Morphotype 2) lateral tooth (29x24x15 mm) (Fanti et al., 2014)
?(MGGCTUN113; Morphotype 2) lateral tooth (27x24x13 mm) (Fanti et al., 2014)
?(MGGCTUN114; Morphotype 2) lateral tooth (48x26x18 mm) (Fanti et al., 2014)
four teeth (Schluter and Schwarzhans, 1978)
Comments-
Fanti et al. (2014) noted "all typical morphological features of
carcharodontosaurid teeth are identifiable in teeth included in
Morphotype 2, including large and moderately curved crowns, strong
labiolingual compression of the basal cross-section, and enamel
wrinkles flanking both serrated carinae", but found "similar
characteristics are observed in the teeth of the abelisaurid[s] Skorpiovenator"
and MGUP MEGA002 from the Duwi Formation of Egypt, so assigned these
"to either a carcharodontosaurid or abelisaurid theropod." They
resolved as abelisaurid when Fanti et al. added them to Hendrickx's
theropod dental matrix, but it would leave carcharodontosaurids
unrepresented in their 68 tooth Dahar sample, which seems
unlikely. The caudal vertebra was assigned to
Carcharodontosauridae by Fanti et al. (2014).
References- Schluter and Schwarzhans, 1978. Eine Bonebed-Lagerstatte
aus dem Wealden Sud-Tunesiens (Umgebung Ksar Krerachfa). Berliner geowissenchaftliche
Abhandlungen A. 8, 53-65.
Bouaziz, Buffetaut, Ghanmi, Jaeger, Martin, Mazin and Tong, 1988. Nouvelles
decouvertes de vertebres fossiles dans l'Albien du Sud tunisien. Bulletin de
la societie geologiques de France, 8th series. 4(2), 335-339.
Benton, Bouaziz, Buffetaut, Martill, Ouaja, Soussi and Trueman, 2000. Dinosaurs
and other fossil vertebrates from fluvial deposits in the Lower Cretaceous of
Southern Tunisia. Palaeogeography, Palaeoclimatology, Palaeoecology. 157, 227-246.
Buffetaut and Ouaja, 2002. A new specimen of Spinosaurus (Dinosauria,
Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the evolutionary
history of the Spinosauridae. Bulletin de la societie geologiques de France.;
173(5), 415-421.
Fanti, Cau, Martinelli and Contessi, 2014. Integrating palaeoecology and morphology
in theropod diversity estimation: A case from the Aptian-Albian of Tunisia.
Palaeogeography, Palaeoclimatology, Palaeoecology. 410, 39-57.
C? sp. indet. (Lapparent, 1951)
Albian-Early Cenomanian, Early Cretaceous-Late Cretaceous
Continental Intercalaire, Tunisia
Material- (MNNHN coll.) five teeth (Lapparent, 1960)
(from Dahar) nine teeth (Lapparent, 1951)
Comments- These teeth are said to have Carcharodontosaurus' 'characteristic
thickness and form', so may be carcharodontosaurid.
References- Lapparent, 1951. Decouverte de dinosauriens associes la und
faunad de reptiles et de poissons, dans le Cretaceous inferieur de l'extreme
sud tunisien. Compte rendu hebdomadaire des seances de l’Academie des Sciences
Paris. 232, 1430-1432.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du
Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
C? sp. indet. (Lapparent, 1960)
Aptian, Early Cretaceous
Elrhaz Formation, Niger
Material- (MNN GDF coll.) postorbital (Taquet, 1976)
?(MNNHN coll.) mid caudal vertebra (>85 mm), radius (~110 mm), manual ungual
(85 mm), pedal phalanx (85 mm) (Lapparent, 1960)
Comments- This material described by Lapparent was not found associated,
so could belong to multiple individuals of several large theropod taxa (Deltadromeus,
Bahariasaurus, Spinosaurus, etc.). Taquet (1976) notes the tooth
MNNHN Td. 2250 is not referrable to Carcharodontosaurus, contra Lapparent.
The tooth fragment MNNHN Td. 2269 described by Lapparent from this locality
is carcharodontosaurid, but Taquet notes it came from a cliff of Albian deposits.
Taquet describes a postorbital as being similar to Acrocanthosaurus,
so this may belong to Carcharodontosaurus.
References- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Taquet, 1976. G�ologie et Pal�ontologie du Gisement de Gadoufaoua
(Aptien du Niger) [Geology and Paleontology of the Gadoufaoua Locality (Aptian
of Niger)]. Cahiers de Pal�ontologie, Centre National de la Recherche
Scientifique, Paris. 191 pp.
C? sp. indet. (Lapparent, 1960)
Berriasian-Barremian, Early Cretaceous
Irhazer Group, Niger
Material- (MNNHN coll.) partial cervical vertebra, two dorsal vertebrae
(120 mm), two sacral vertebrae (280 mm combined), mid caudal vertebra (115 mm),
three caudal vertebrae (100, 110, 120 mm), two distal chevrons (Lapparent, 1960)
Comments- This material was not found associated, so could belong to
multiple individuals of several large theropod taxa (Deltadromeus, Bahariasaurus,
Spinosaurus, etc.).
Reference- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
C? sp. indet. (Lapparent, 1960)
Early Cretaceous
Continental Intercalaire, Niger
Material- (MNNHN coll.; from Tefidet) two mid caudal vertebrae (85 mm)
(Lapparent, 1960)
(from Akarazeras) teeth (Taquet, 1976)
Comments- This material was not found associated, so could belong to
multiple individuals of several large theropod taxa (Deltadromeus, Bahariasaurus,
Spinosaurus, etc.).
References- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Taquet, 1976. G�ologie et Pal�ontologie du Gisement de Gadoufaoua
(Aptien du Niger) [Geology and Paleontology of the Gadoufaoua Locality (Aptian
of Niger)]. Cahiers de Pal�ontologie, Centre National de la Recherche
Scientifique, Paris. 191 pp.
C? sp. indet. (Lapparent, 1960)
Early Cretaceous
Continental Intercalaire, Sahara Desert
Material- (MNNHN coll.) eight vertebrae, partial humerus, distal manual
phalanx (Lapparent, 1960)
Comments- This material was not found associated, so could belong to
multiple individuals of several large theropod taxa (Deltadromeus, Bahariasaurus,
Spinosaurus, etc.).
Reference- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
C? sp. indet. (Buffetaut, 1989)
Late Albian-Early Cenomanian, Early Cretaceous
"Red Beds" (Tegama Formation?), Hammada du Guir, Morocco
Material- (IMGP coll.) teeth (Buffetaut, 1989)
Late Albian-Early Cenomanian, Early Cretaceous
"Red Beds", Ouaouizaght, Morocco
material (Allain and Aquesbi, 2008)
Comments- Buffetaut (1989) mentioned "the compressed and serrated teeth of a "normal" carnivorous dinosaur, probably Carcharodontosaurus
Stromer." Allain and Aquesbi (2008) noted "the Albian-Cenomanian
red deposits have yielded carcharodontosaurid remains in the vicinity
of Ouaouizaght (pers. obs.)."
References- Buffetaut, 1989. New remains of the enigmatic dinosaur Spinosaurus
from the Cretaceous of Morocco and the affinities between Spinosaurus
and Baryonyx. Neues Jahrbuch f�r Geologie und Pal�ontologie
Monatshefte. 1989(2), 79-87.
Allain and Aquesbi, 2008. Anatomy and phylogenetic relationships of Tazoudasaurus naimi (Dinosauria, Sauropoda) from the late Early Jurassic of Morocco. Geodiversitas. 30(2), 345-424.
C? sp. indet. (Lapparent, 1960)
Albian, Early Cretaceous
Continental Intercalaire, Algeria
Material- ?(MNNHN coll.; from Alrar) caudal vertebra (60 mm) (Lapparent,
1960)
(MNNHN coll.; from Aoulef) dorsal centrum (75 mm), proximal caudal centrum (105
mm), few mid caudal vertebrae (~70-100 mm), caudal vertebra (80 mm) (Lapparent,
1960)
(from Oued Boudjihane) teeth (Bassoullet and Iliou, 1967)
Comments- This material was not found associated, so could belong to
multiple individuals of several large theropod taxa (Deltadromeus, Bahariasaurus,
Spinosaurus, etc.). A manual ungual from Dijoua referred to C. saharicus
by Lapparent matches bone taxon J of Russell (1996), identified as Spinosaurus
by Ibrahim et al. (2014). A distal metatarsal from Alrar referred to C. saharicus
by Lapparent is actually a manual phalanx and matches bone taxon I of Russell.
Russell notes it could belong to the same taxon as the ungual, so this may be
Spinosaurus as well. Finally, a pedal ungual from Alrar referred to C.
saharicus by Lapparent matches the Spinosaurus B morphotype which
has also been identified as Spinosaurus by Ibrahim et al..
Bassoullet and Iliou (1967) reported (translated) "numerous teeth of carnivorous theropods
some of which are attributable to the genus Carcharodontosaurus, others to coelurosaurids" and "1
vertebra (probably cervical) of a large theropod." Bassoullet worked
at the Facult� des sciences de Paris, but this was dissolved in 1970 so
that if the remains were stored there they may have been moved to one
of the University of Paris campuses or the University of Orleans.
References- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Bassoullet and Iliou, 1967. Decouverte de Dinosaures associes a des
Crocodiliens et des Poissons dans le Cretace inferieur de l'Atlas
saharien (Algerie). Compte rendu sommaire des s�ances de la Soci�t�
g�ologique de France. 7, 294-295.
Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt,
Morocco. Bulletin du Muse'um national d'Histoire naturelle (4e se'r.). 18, 349-402.
Ibrahim, Sereno, Dal Sasso, Maganuco, Fabbri, Martill, Zouhri, Myhrvold and
Iurino, 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science.
345(6204), 1613-1616.
C? sp. indet. (Bond and Bromley, 1970)
Late Jurassic or Early Cretaceous
Gokwe Formation, Zimbabwe
Material- teeth
Reference- Bond and Bromley, 1970. Sediments with the remains of dinosaurs
near Gokwe, Rhodesia. Palaeogeography, Palaeoclimatology, Palaeoecology. 8,
313-327.
C? sp. indet. (Churcher, 1995)
Early-Middle Campanian, Late Cretaceous
El Atrun, Quseir Formation, Kharga Oasis, Egypt
Material- teeth (Churcher, 1995)
Early-Middle Campanian, Late Cretaceous
Bee's Friday Site, Quseir Formation, Dakhla Oasis, Egypt
(ROM? coll.) teeth and/or bones (Churcher, 1999)
Comments- According to Churcher (1995), "On a visit to the El Atrun exposure in 1993, Dr. Dale A. Russell of the Canadian Museum of Nature, Ottawa, and I found isolated teeth of two carnivorous dinosaurs, tentatively assigned to Spinosaurus and Carcharodontosaurus...", but this to too late to be congeneric with the Cenomanian Carcharodontosaurus.
Churcher
and Russell (1992) stated in the Dakhla (spelled Dakhleh by them)
Oasis, "theropod and sauropod teeth and bones were found a few metres
below the Duwi Formation." Churcher's (1999) faunal list matches this
with Bee's Friday Site, and says "Subsequently specimens assignable to Spinosaurus and Carcharodontosaurus were obtained", suggesting this was the theropod material mentioned in 1992.
References-Churcher and Russell, 1992. Terrestrial vertebrates from Campanian
strata in Wadi el-Gedid (Kharga and Dakhleh Oases), Western Desert of
Egypt. Journal of Vertebrate Paleontology. 12(3), 23A.
Churcher, 1995. Giant Cretaceous lungfish Neoceratodus tuberculatus
from a deltaic environment in the Quseir (=Baris) Formation of Kharga
oasis, Western Desert of Egypt. Journal of Vertebrate Paleontology.
15(4), 845-849
Churcher,
1999. A note on the Late Cretaceous vertebrate fauna of the Dakhleh
Oasis. In Churcher and Mills (eds.). Reports from the Survey of Dakhleh
Oasis, Western Desert of Egypt, 1977-1987. Dakhleh Oasis Project:
Monograph 2. Oxbow Monograph 99. 55-67..
Giganotosaurinae Coria and Currie, 2006
Definition- (Giganotosaurus carolinii, Mapusaurus rosea
<- Carcharodontosaurus saharicus) (modified from Coria and Currie,
2006)
= Giganotosaurini Coria and Currie, 2006 vide Brusatte and Sereno, 2008
Definition- (Giganotosaurus carolinii <- Carcharodontosaurus saharicus)
(Brusatte and Sereno, 2008)
Diagnosis- (after Coria and Currie, 2006) femur with a weak fourth trochanter;
shallow and broad extensor groove.
(proposed) short distal caudal prezygapophyses (<25% of central length); humerus with broad distal end and little separation
between condyles; cruciate ridge in femoral flexor groove absent.
References- Coria and Currie, 2006. A new carcharodontosaurid (Dinosauria,
Theropoda) from the Upper Cretaceous of Argentina. Geodiversitas. 28(1), 71-118.
Meraxes Canale, Apestegu�a, Gallina, Mitchell, Smith, Cullen, Shinya, Haluza, Gianechini and Makovicky, 2022
M. gigas Canale, Apestegu�a, Gallina, Mitchell, Smith, Cullen, Shinya, Haluza, Gianechini and Makovicky, 2022
Late Cenomanian, Late Cretaceous
La Campanas Canyon, Huincul Formation of Rio Limay Subgroup, Neuquen, Argentina
Holotype- (MMCh-PV 65; Campanas carcharodontosaurid) (~10.7
m, ~4.3 tons; ~40 year old adult) incomplete skull (~1.27 m), partial
~fourth cervical vertebra (160 mm), four cervical neural spines,
posterior dorsal centrum (154 mm), dorsal neural spine fragment, six
dorsal rib fragments, gastralia, synsacrum, first-fifteenth caudal
vertebrae (c1 182, c5 158, c10 146, c15 138 mm), first-fifteenth
chevrons, scapulae (790 mm), incomplete coracoids, humeri (340 mm),
radii (186 mm), ulnae (228 mm), intermedium, distal carpal I, phalanx
I-1 (93 mm), manual unguals I (60 mm), metacarpals II (85 mm),
phalanges II-1 (86 mm), incomplete phalanx II-2, manual ungual II (46
mm), metacarpal III (65 mm), phalanx III-1 (34 mm), manual ungual III
(32 mm), ilia (1.080 m), pubes (1.090 m), ischia (1.015 m), femora
(1.260 m), tibiae (1.090 m), fibulae (940 mm), astragali (224 mm
trans), calcanea, distal tarsal IV, metatarsal I (94 mm), pedal unguals
I (76 mm), metatarsal II (436 mm), phalanges II-1, phalanges II-2,
pedal ungual II, metatarsal III (486 mm), phalanges III-1, phalanges
III-2, phalanges III-3, pedal unguals III, metatarsals IV (443 mm),
phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal
ungual IV, metatarsals V (192 mm)
Diagnosis- (after Canale et al., 2022) maxillary fenestra; jugal stepped along posterior edge of postorbital process (unknown in Giganotosaurus);
postorbital with a low and rounded lateral horn on the posterior
process; lacrimal with rounded projections along its dorsal margin
(unknown in Mapusaurus? and Tyrannotitan); sacral vertebral neural spines almost completely co-ossified (unknown in Mapusaurus; ontogenetic?); caudal vertebrae 1-4 with hyposphene-hypantrum articulations (unknown in Mapusaurus and Tyrannotitan);
humerus with internal tuberosity separated from humeral head by a cleft
(unknown in other giganotosaurines); humerus with ovoid fossa on its
anteroproximal surface (unknown in other giganotosaurines); strongly
enlarged pedal ungual II, almost twice the length of pedal ungual IV
(unknown in other giganotosaurines).
Other diagnoses- In addition to
the diagnostic postorbital and jugal caharcters retained
in its eventual description, Canale et al. (2018) also cite "proximal
caudal vertebrae with extensive pneumatic openings in the bases of the
neural spines above the postzygapophyses." The description
indicates these are "wide and deep spinopostzygapophyseal fossae along
the posterior base of the neural spines, in the first five" and that "A
pair of foramina open into these fossae behind the
postzygapophyses." As with the proximal caudal
hyposphene-hypantra, preserved caudals of Mapusaurus and Tyrannotitan are more distally positioned and while Giganotosaurus might be assumed to lack them this remains undescribed.
Canale et al. (2022) also list "quadratojugal with a deep and rounded
fossa on its lateral surface", but this isn't visible in their figure
2A and the element is unpreserved in Mapusaurus and Giganotosaurus in any case.
Comments- Discovered in 2012
and excavated from 2012 to 2014, this was first mentioned by Canale et
al. (2018) in an abstract as "A recently recovered specimen with
carcharodontosaurid affinities (MMCh-PV 65) from the Huincul Formation
(Cenomanian)." They state some features "provide evidence for a
possible new taxon" and that "A preliminary phylogenetic analysis
recovers the specimen nested within Giganotosaurini." Cullen et
al. (2020) call this the "Campanas carcharodontosaurid (unnamed
carcharodontosaurid taxon from the Huincul Formation of Argentina; MMCh
PV 65)" and figure sections of a rib, gastralium, femur and
fibula. Palombi (2022) described the metatarsals in his thesis,
calling it "the holotype of a new species of Carcharodontosauridae
(MMCh-PV 65), which is currently under study" (translated).
Canale et al. (2022) described the specimen as Meraxes gigas,
and using Canale's carnosaur analysis recovered it as the basalmost
giganotosaurine. Canale (pers. comm., 7-2002) confirms that in
the right hindlimb "we have fibula, calcaneum, metatarsals IV and V.
The digits of the right foot are incomplete, just some phalanges (five
preungual, two unguals)", which is not determinable in the publication.
References- Canale, Apesteguia,
Makovicky, Gallina, Smith, Mitchell, Gianechini and Haluza, 2018. New
Carcharodontosauridae (Dinosauria, Theropoda) from the early Late
Cretaceous of Neuquen, Argentina, yields light on the anatomy of the
group. Reuni�n de Comunicaciones de la Asociaci�n Paleontol�gica
Argentina, Libro de Resumenes. R14.
Cullen, Canale, Apestegu�a, Smith, Hu and Makovicky, 2020.
Osteohistological analyses reveal diverse strategies of theropod
dinosaur body-size evolution. Proceedings of the Royal Society B.
287(1939), 20202258.
Canale, Apestegu�a, Gallina, Mitchell, Smith, Cullen, Shinya, Haluza,
Gianechini and Makovicky, 2022. New giant carnivorous
dinosaur reveals convergent evolutionary trends in theropod arm
reduction. Current Biology. 32(14), 3195-3202.
Palombi, 2022. Osteolog�a y miolog�a del autopodio posterior en un
nuevo dinosaurio Carcharodontosauridae de la Formaci�n Huincul
(Cenomaniano-Turoniano, Provincia de Neuqu�n, Argentina). Licenciado en
Paleontolog�a thesis, Universidad Nacional de R�o Negro. 69 pp.
Tyrannotitan Novas, de
Valais, Vickers-Rich and Rich, 2005
T. chubutensis Novas, de Valais, Vickers-Rich and Rich, 2005
Albian, Early Cretaceous
La Juanita farm, Cerro Casta�o Member of the Cerro Barcino Formation, Chubut, Argentina
Holotype- (MPEF-PV 1156) (~11.4 m) incomplete dentaries, partial second
dorsal neural arch, partial third dorsal vertebra, incomplete fourth dorsal
vertebra, incomplete fifth dorsal vertebra, incomplete sixth dorsal vertebra,
seventh dorsal vertebra, eighth dorsal neural spine, ninth dorsal neural spine,
tenth dorsal vertebra, eleventh dorsal vertebra, fourteenth dorsal neural spine,
gastral fragments, first sacral neural spine, proximal caudal vertebra, six
chevrons, proximal scapulocoracoid, distal humerus, radii (one partial, one
fragmentary), ilial fragments, incomplete pubes, incomplete ischia, femora,
fibula
Paratype- (MPEF-PV 1157) (~12.2 m) incomplete jugal, quadratojugal, incomplete
dentary (680 mm), two teeth, atlas, incomplete seventh cervical vertebra, incomplete
first dorsal vertebra, incomplete fourth dorsal vertebra, incomplete sixth dorsal
vertebra, seventh dorsal vertebra, partial eighth dorsal neural arch, twelfth
dorsal centrum, thirteenth dorsal centrum, fourteenth dorsal centrum, dorsal
rib fragment, fourteenth dorsal rib, partial sacrum, incomplete distal caudal
vertebra, chevron, femur (1.40 m), distal metatarsal II, phalanx II-2, phalanx IV-2, phalanx IV-3, pedal
ungual ?IV
Referred- (MPEF-PV 10821) nineteen teeth (Canale et al., 2015)
Diagnosis- (after Novas et al., 2005) some teeth with bilobate denticles on
mesial carina (unknown in Meraxes).
(after Canale et al., 2015) dentary with an anteroventrally-posterodorsally
symphyseal margin in lateral view (unknown in Meraxes); second and third dorsal vertebrae with well-developed
accessory lamina connecting anterior and posterior centrodiapophyseal laminae (unknown in Meraxes);
fibular fossa extended far proximal to ectocondylar tuber on the distal
femur; anterior margin of proximomedial fibular fossa posteriorly
projected, covering part of the fossa.
Other diagnoses- Canale et al. (2015) noted the depth of the Meckelian
groove is similar to Carcharodontosaurus? iguidensis, Giganotosaurus
and Allosaurus, thus Novas et al.'s (2005) deep Meckelian groove is invalid.
Similarly, Canale et al. noted the posterior dorsal neural spine ligament scars
are equally well developed in Acrocanthosaurus, Giganotosaurus
and Mapusaurus, contra Novas et al.. Note both hindlimb characters listed above should be scorable in Meraxes but are not determinable in its description.
Comments-
The material was discovered in 1994 (Canale, pers. comm. 7-2022), with the holotype
and paratype about 1 km apart. Rich et al. (2000) preliminarily
described both as unnamed theropods, noting MPEF-PV 1157 may be
conspecific as "suggested by the presence of a bone fragment which
appears to be a heavily worn specimen of a neural spine with triangular
bosses like those on the neural spines of MPEF-PV 1156." They
listed "a possible maxilla fragment" and "a partial ilium" in the
material of MPEF-PV 1157, which must have been misidentified.
Novas et al. (2005) list three pedal phalanges as being present in the
holotype ("2.I, 2.II and 3.III"), but also illustrate an ungual
supposedly from digit II, which wouldn't correspond to any of these.
The skeletal reconstruction only shows two phalanges- ungual III and a
distal phalanx. Canale et al. (2015) reidentified the phalanges as
II-2, ungual II, IV-2 and IV-3. The supposed ulna was reidentified as a
radius and several presacral centra had revised positions in Canale et
al.. Based on comparison to Meraxes
and the small size of pedal ungual II compared to phalanx II-2 in
Canale et al.'s Figure 33, the pedal ungual is reassigned to digit IV.
Note MPEF-PV 10821 are "isolated teeth found in this locality, which
are housed at the MPEF collections but have not been assigned to the
holotype or paratype" (Canale et al., 2015).
References- Rich, Vickers-Rich, Novas, C�neo, Puerta and Vacca,
2000. Theropods from the "Middle" Cretaceous Chubut Group of the San
Jorge sedimentary basin, central Patagonia. A preliminary note. GAIA. 15, 111-115.
Novas, de Valais, Vickers-Rich and Rich, 2005. A large Cretaceous theropod from
Patagonia, Argentina, and the evolution of carcharodontosaurids. Naturwissenschaften.
92(5), 226-230.
Canale, 2010. Los Carcharodontosauridae (Dinosauria, Theropoda) del
Cretacico de America del Sur: Anatomıa, relaciones filogeneticas y
paleobiologıa. PhD Thesis, Universidad Nacional de La Plata. 261 pp.
Gianechini, Pol and Vieytes, 2011. Enamel microstructure of theropod
teeth: A characterization of specimens from the Cretaceous of
Patagonia. Ameghiniana. 48(4), R167-R168.
Canale, Novas and Pol, 2015. Osteology and phylogenetic relationships of Tyrannotitan
chubutensis Novas, de Valais, Vickers-Rich and Rich, 2005 (Theropoda: Carcharodontosauridae)
from the Lower Cretaceous of Patagonia, Argentina. Historical Biology. 27(1),
1-32.
Mapusaurus Coria and Currie, 2006
= "Mapusaurus" Fiorillo and Eberth, 2004
= Taurovenator Motta, Aranciaga
Rolando, Rozadilla, Agnolin, Chimento, Brisson Egli and Novas, 2016
M. roseae Coria and Currie, 2006
= "Mapusaurus rosae" Papolio, 2004
= Taurovenator violantei Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento,
Brisson Egli and Novas, 2016
Late Cenomanian, Late Cretaceous
Ca�ad�n del Gato, Huincul Formation of Rio Limay Subgroup, Neuquen, Argentina
Holotype- (MCF-PVPH-108.1) nasal
Paratypes- (MCF-PVPH-108.5) incomplete lacrimal/prefrontal
(MCF-PVPH-108.45) incomplete humerus (~300 mm)
(MCF-PVPH-108.83) axial neural arch
(MCF-PVPH-108.90) mid cervical neural arch
(MCF-PVPH-108.115) maxilla
(MCF-PVPH-108.125) partial dentary
(MCF-PVPH-108.128) ilium (1.05 m)
(MCF-PVPH-108.165) ischium (1.01 m)
(MCF-PVPH-108.167) incomplete jugal
(MCF-PVPH-108.177) postorbital
(MCF-PVPH-108.179) splenial
(MCF-PVPH-108.202) (~12.6 m) fibula (860 mm)
Referred- (MCF-PVPH-108.2) partial dentary (Coria and Currie, 2006)
(MCF-PVPH-108.3) (~5.5 m; juvenile) partial dentary (Coria and Currie, 2006)
(MCF-PVPH-108.4) postorbital (Coria and Currie, 2006)
(MCF-PVPH-108.6) quadrate (Coria and Currie, 2006)
(MCF-PVPH-108.7) angular fragment (Coria and Currie, 2006)
(MCF-PVPH-108.8) anterior dentary tooth (65x33x20 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.9) mid dentary tooth (71x32x17 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.10) posterior tooth (41x25x13 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.11) maxillary fragment (Coria and Currie, 2006)
(MCF-PVPH-108.12) anterior nasal fragment (Coria and Currie, 2006)
(MCF-PVPH-108.14) manual ungual II(?) (Coria and Currie, 2006)
(MCF-PVPH-108.15) partial surangular (Coria and Currie, 2006)
(MCF-PVPH-108.16) tooth (50x28x15 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.17) posterior nasal fragment (Coria and Currie, 2006)
(MCF-PVPH-108.18) pedal phalanx IV-2 (Coria and Currie, 2006)
(MCF-PVPH-108.19) pedal phalanx IV-1 (Coria and Currie, 2006)
(MCF-PVPH-108.21) pedal phalanx IV-2 (Coria and Currie, 2006)
(MCF-PVPH-108.22) pedal phalanx IV-2 (Coria and Currie, 2006)
(MCF-PVPH-108.23) pedal phalanx III-1 (Coria and Currie, 2006)
(MCF-PVPH-108.24) pedal phalanx III-2 (Coria and Currie, 2006)
(MCF-PVPH-108.25) partial femur, pedal phalanx III-2 (Coria and Currie, 2006)
(MCF-PVPH-108.26) pedal phalanx III-1 (Coria and Currie, 2006)
(MCF-PVPH-108.27) pedal phalanx II-2(?) (Coria and Currie, 2006)
(MCF-PVPH-108.28) pedal phalanx III-3 (Coria and Currie, 2006)
(MCF-PVPH-108.31) (~6.4 m) metatarsal III (454 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.32, 34-36) (~6.0-6.1 m) metatarsal II, metatarsal II (385 mm),
metatarsal III (434 mm), metatarsal IV (Coria and Currie, 2006)
(MCF-PVPH-108.33, 188) (~6.5-7.2 m) metatarsal II (450 mm), metatarsal III (460
mm) (Coria and Currie, 2006)
(MCF-PVPH-108.37) (~7.3 m) metatarsal IV (475 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.38, 200) (~6.6 m) metatarsal II (415 mm), metatarsal III (>410
mm) (Coria and Currie, 2006)
(MCF-PVPH-108.39) partial dentary (Coria and Currie, 2006)
(MCF-PVPH-108.41) tooth (FABL >23 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.42) tooth (33x17.7x13.5 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.43) tooth (53x31x14.5 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.44) (~9.9 m) femur (Coria and Currie, 2006)
(MCF-PVPH-108.46) radius (Coria and Currie, 2006)
?(MCF-PVPH-108.48) fused proximal metacarpal II and III (or distal humerus)
(Coria and Currie, 2006)
(MCF-PVPH-108.50) scapula (Coria and Currie, 2006)
(MCF-PVPH-108.51) partial fibula (Coria and Currie, 2006)
(MCF-PVPH-108.52) partial tibia (Coria and Currie, 2006)
(MCF-PVPH-108.53) partial tibia (Coria and Currie, 2006)
(MCF-PVPH-108.54) partial femur (Coria and Currie, 2006)
(MCF-PVPH-108.57) partial femur (Coria and Currie, 2006)
(MCF-PVPH-108.58) (~9.7 m) tibia (Coria and Currie, 2006)
(MCF-PVPH-108.59) partial femur (Coria and Currie, 2006)
(MCF-PVPH-108.61) partial femur (Coria and Currie, 2006)
(MCF-PVPH-108.62) partial tibia (Coria and Currie, 2006)
(MCF-PVPH-108.63) partial tibia (Coria and Currie, 2006)
(MCF-PVPH-108.64) partial femur (Coria and Currie, 2006)
(MCF-PVPH-108.65) partial femur (Coria and Currie, 2006)
(MCF-PVPH-108.66) partial tibia (Coria and Currie, 2006)
(MCF-PVPH-108.67) (~8.1 m) tibia (Coria and Currie, 2006)
(MCF-PVPH-108.68) (~9.8 m) tibia (1.04 m) (Coria and Currie, 2006)
(MCF-PVPH-108.69) scapular fragment (Coria and Currie, 2006)
(MCF-PVPH-108.70) incomplete astragalus (Coria and Currie, 2006)
(MCF-PVPH-108.71) partial coracoid (Coria and Currie, 2006)
(MCF-PVPH-108.73) partial tibia (Coria and Currie, 2006)
(MCF-PVPH-108.75) mid caudal vertebra (Coria and Currie, 2006)
(MCF-PVPH-108.76) mid caudal vertebra (165 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.78) mid caudal vertebra (Coria and Currie, 2006)
(MCF-PVPH-108.79) distal caudal vertebra (97 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.80) posterior dorsal centrum (165 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.81) proximal caudal vertebra (140 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.82) anterior dorsal vertebra (87 mm; excluding anterior ball)
(Coria and Currie, 2006)
(MCF-PVPH-108.84) mid dorsal neural arch (Coria and Currie, 2006)
(MCF-PVPH-108.85) dorsal neural arch (Coria and Currie, 2006)
(MCF-PVPH-108.89) fifth sacral centrum (135 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.95) proximal ischium (Coria and Currie, 2006)
(MCF-PVPH-108.96) proximal ischium (Coria and Currie, 2006)
(MCF-PVPH-108.97) dorsal chevron (Coria and Currie, 2006)
(MCF-PVPH-108.100) lacrimal (Coria and Currie, 2006)
(MCF-PVPH-108.101) lacrimal (Coria and Currie, 2006)
(MCF-PVPH-108.102) quadrate (Coria and Currie, 2006)
(MCF-PVPH-108.103) posterior dentary tooth (24x20x9 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.106) anterior dorsal rib (Coria and Currie, 2006)
(MCF-PVPH-108.109) manual phalanx II-2 (80 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.110) tooth (81.5x30x10.5 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.111) tooth (77x38x17 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.113) tooth (54x19x8.5 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.114) tooth (Coria and Currie, 2006)
(MCF-PVPH-108.116) furcula or anterior gastralium (Coria and Currie, 2006)
(MCF-PVPH-108.120) tooth (36x22 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.124) metatarsal II (Coria and Currie, 2006)
(MCF-PVPH-108.131) tooth (?x19x8 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.132) (~8.4 m) fibula (Coria and Currie, 2006)
(MCF-PVPH-108.138) maxilla, tooth (47x23 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.139) partial prearticular (Coria and Currie, 2006)
(MCF-PVPH-108.141) tooth (39x28x12 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.142) maxilla (Coria and Currie, 2006)
(MCF-PVPH-108.145) (~12.2 m) pubic shaft (Coria and Currie, 2006)
(MCF-PVPH-108.148) proximal pubis (Coria and Currie, 2006)
(MCF-PVPH-108.149) proximal pubis (Coria and Currie, 2006)
(MCF-PVPH-108.153) postorbital (Coria and Currie, 2006)
(MCF-PVPH-108.162) cervical epipophysis (Coria and Currie, 2006)
(MCF-PVPH-108.166) tooth (42x23x16 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.168) jugal (Coria and Currie, 2006)
(MCF-PVPH-108.169) partial maxilla, tooth (68 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.170) quadrate (Coria and Currie, 2006)
(MCF-PVPH-108.171) tooth (56x29x16 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.173) tooth (>73x37 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.175) partial dorsal rib (Bell and Coria, 2013)
(MCF-PVPH-108.176) tooth (Coria and Currie, 2006)
(MCF-PVPH-108.180) tooth (Coria and Currie, 2006)
(MCF-PVPH-108.181) ilial fragment (>1.05 m) (Coria and Currie, 2006)
(MCF-PVPH-108.183) incomplete lacrimal/prefrontal (Coria and Currie, 2006)
(MCF-PVPH-108.185) (~12.2 m) scapular fragment (Coria and Currie, 2006)
(MCF-PVPH-108.187) scapular fragment (Coria and Currie, 2006)
(MCF-PVPH-108.189) (~8.3 m) fibula (Coria and Currie, 2006)
(MCF-PVPH-108.196) fibula (Coria and Currie, 2006)
(MCF-PVPH-108.198) pedal ungual ?IV (Coria and Currie, 2006)
(MCF-PVPH-108.201) (~6.3 m) metatarsal III (450 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.203) (~10.2 m) femur (Coria and Currie, 2006)
(MCF-PVPH-108.205) mid caudal vertebra (120 mm) (Coria and Currie, 2006)
(MCF-PVPH-108.209) first sacral centrum (Coria and Currie, 2006)
(MCF-PVPH-108.210) dorsal chevron (Coria and Currie, 2006)
(MCF-PVPH-108.220) partial fibula (Coria and Currie, 2006)
(MCF-PVPH-108.230) gastralium fragment (Coria and Currie, 2006)
(MCF-PVPH-108.233) (~9.5 m) femur (Coria and Currie, 2006)
(MCF-PVPH-108.234) partial femur (1.30 m) (Coria and Currie, 2006)
(MCF-PVPH-108.245) partial ilium (~1.05 m) (Coria and Currie, 2006)
(MCF-PVPH-108.246) metatarsal I (Coria and Currie, 2006)
(MCF-PVPH-108.247) distal caudal vertebra (44 mm) (Coria and Currie, 2006)
(MCF-PVPH-108 coll.) few dorsal ribs, hundreds of dorsal rib fragments, gastralium
fragments, more than nine pedal phalanges (Coria and Currie, 2006)
Late Cenomanian, Late Cretaceous
Violante farm, Huincul Formation of Rio Limay Subgroup, Rio Negro, Argentina
(MPCA PV 802; holotype of Taurovenator violantei) postorbital (Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016)
Diagnosis- (after Coria and Currie, 2006) upper quadratojugal process
of jugal splits into two prongs (unknown in Giganotosaurus);
anterior mylohyoid foramen positioned above dentary contact on splenial
(unknown in other giganotosaurines); second and third metacarpals fused
(supposed carpometacarpus may be distal humerus; unknown in Giganotosaurus and Tyrannotitan); brevis fossa of ilium extends
deeply into excavation dorsal to ischial peduncle.
Other diagnoses- Coria and Currie (2006) listed "humerus with broad distal end and little separation
between condyles", but this has since been found in Meraxes and is unpreserved in other giganotosaurines.
Comments- The type material comes from at least nine individuals. Carrano
et al. (2012) believe MCF-PVPH-108.48 is a distal humerus and not a carpometacarpus
as Coria and Currie (2006) described it. While non-maniraptoriforms otherwise
lack metacarpal fusion and it does resemble a distal humerus, it differs from
Mapusaurus
in having two bulbous condyles with a notch between them and a large
epicondyle, so may belong to another theropod. MCF-PVPH-108.220 was
described as a partial fibula by Coria and Currie, but reidentified as
a pathological dorsal rib by Bell and Coria (2013). Coria and
Currie state the preserved pedal ungual (MCF-PVPH-108.198) "is
asymmetrical and was probably from either the second or fourth digits",
and based on its shortness and high degree of curvature it is here
identified as ungual IV using the complete pes of Meraxes for comparison.
This taxon was discovered in 1995, but only reported to Coria in 1997, when
he and Currie examined the material. It was announced at that years Society
of Vertebrate Paleontology meeting, and described briefly in an abstract (Coria
and Currie, 1997). At the time, only the remains of an 8 meter long specimen
were known, and it was identified as an adult. Coria and Currie returned to
the site in 1998 to discover the presence of at least six individuals, some
of which Currie said could be larger than Giganotosaurus’ holotype.
The largest specimens are MCF-PVPH-145, 185 and 202, which are about 100-103%
the size of the Giganotosaurus holotype.
The association of several individuals was suggested to be due to pack behavior.
This was reported to the popular media in May 1999, and later described in another
abstract (Eberth et al., 2000). Later (Eberth and McCrea, 2001), the minimum
number of individuals was increased to eight. This paper finds the probable
cause of death to be drought and notes the bones experienced at least two flooding
events and were exposed and trampled over more than one season. However, they
state several alternatives exist besides gregarious behavior to explain the
find, including environmental stress and breeding. In the final publication,
Coria and Currie (2006) raised the minimum number of individuals to nine.
It was reported on the internet that a magazine had termed the taxon Giganotosaurus
"argentine", but this has yet to be confirmed and would be a nomen
nudum in any case. Fiorillo and Eberth (2004) used the name "Mapusaurus"
in their taphonomy chapter in The Dinosauria 2nd. edition, probably by accident.
Papolio (2004) listed it as "Mapusaurus rosae" in a field guide.
Motta et al. (2016) described the postorbital MPCA PV 802 as a new carcharodontosauirine Taurovenator violantei. They diagnosed it based on two characters- "horn-like prominence on orbital
brow of postorbital" is present but less developed in Mapusaurus
lacrimal MCF-PVPH-108.177, as noted by Coria et al. (2019); while "deep
excavation in ventral surface of postorbital" was stated to be unknown
in Mapusaurus as "the region
in which the fossa should be housed is broken." Yet the latter
seems to be illustrated as present by Coria and Currie (2006: Fig. 5D),
and indeed Coria et al. state it is present in Mapusaurus and Canale et al. (2022) find it is also present in Meraxes. Unsurprisingly, Coria et al. suggest Taurovenator is a junior synonym of Mapusaurus, also noting they share a character Coria and Currie lists as distinguishing the latter from Giganotosaurus ("ventrolaterally curving lateral margin of palpebral") although this was later revealed to be present in Meraxes as well. While Taurovenator's
type is smaller than MCF-PVPH-108.177 so might be expected to have a
less developed postorbital horn, interspecific variation in cranial
excrescences is common in theropods (e.g. Allosaurus, Tyrannosaurus). Thus I provisionally agree with the synonymy.
References- Coria and Currie, 1997. A new theropod from the Rio Limay
Formation. Journal of Vertebrate Paleontology. 17(3), 40A.
Eberth, Currie, Coria, Garrido and Zonneveld, 2000. Large-theropod bonebed,
Neuquen, Argentina: Paleoecological importance. Journal of Vertebrate Paleontology.
20(3), 39A.
Eberth and Crea, 2001. Were large theropods gregarious? Journal of Vertebrate
Paleontology. 21(3), 46A-47A.
Fiorillo and Eberth, 2004. Dinosaur taphonomy. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria: Second Edition. 607-613.
Papolio, 2004. Animales Prehist�ricos de Am�rica del Sur. Carlos
Papolio. 96 pp.
Coria and Currie, 2006. A new carcharodontosaurid (Dinosauria, Theropoda) from
the Upper Cretaceous of Argentina. Geodiversitas. 28(1), 71-118.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Bell and Coria, 2013. Palaeopathological survey of a population of Mapusaurus
(Theropoda: Carcharodontosauridae) from the Late Cretaceous Huincul Formation,
Argentina. PLoS ONE. 8(5), e63409.
Canale, Novas, Salgado and Coria, 2015. Cranial ontogenetic variation in Mapusaurus
roseae (Dinosauria: Theropoda) and the probable role of heterochrony in
carcharodontosaurid evolution. Pal�ontologische Zeitschrift. 89(4), 983-993.
Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016. New theropod fauna from the Upper Cretaceous (Huincul
Formtation) of northwestern Patagonia, Argentina. In Khosla and Lucas (eds.).
Cretaceous period: Biotic diversity and biogeography. New Mexico Museum of Natural
History and Science Bulletin. 71, 231-253.
Coria, Currie, Ortega and Baiano, 2019. An Early Cretaceous,
medium-sized carcharodontosaurid theropod (Dinosauria, Saurischia) from
the Mulichinco Formation (upper Valanginian), Neuqu�n Province,
Patagonia, Argentina. Cretaceous Research. Journal Pre-proof DOI:
10.1016/j.cretres.2019.104319
Canale, Apestegu�a, Gallina, Mitchell, Smith, Cullen, Shinya, Haluza,
Gianechini and Makovicky, 2022. New giant carnivorous dinosaur
reveals convergent evolutionary trends in theropod arm reduction.
Current Biology. 32(14), 3195-3202.
Giganotosaurus Coria and
Salgado, 1995
G. carolinii Coria and Salgado, 1995
= Carcharodontosaurus carolinii (Coria and Salgado, 1995) Figueiredo,
1998
Early Cenomanian, Late Cretaceous
Candeleros Formation of Rio Limay Subgroup, Neuquen, Argentina
Holotype-
(MUCPv-Ch1) (12.2 m, 5 tons) (skull ~1.62 m) premaxilla, maxilla,
maxillary teeth (74, 97 mm), nasal, lacrimal, postorbital, quadrates
(410 mm), braincase, ectopterygoid, pterygoid, anterior dentary,
dentary tooth, tooth (>82x45x18 mm), tooth (102x39.5x22 mm), tooth
(88x33.5x20 mm), axis, nine cervical vertebrae, ten dorsal vertebrae,
dorsal ribs, sacrum, eighteen(+?) caudal vertebrae, chevrons, partial
scapulocoracoid, ilium (~1.54 m), pubes (~1.11 m), ischia (~1.2 m), femora
(1.43 m), tibia (1.11 m), fibula (840 mm)
Referred- (MUCPv-52) tooth (90x45x21 mm) (Calvo, 1999)
(MUCPv-95) (~13.2 m, 6.2 tons) (skull ~1.75 m) incomplete dentary, teeth (Calvo, 1989)
Early Cenomanian, Late Cretaceous
La Buitrera, Candeleros Formation of Rio Limay Subgroup, Rio Negro, Argentina
(MPCA coll.) maxillary tooth (99x42x18 mm) (Valais and Apesteguia, 2001)
Early Cenomanian, Late Cretaceous
? Candeleros Formation of Rio Limay Subgroup, Argentina
(FPDM uncatalogued) tooth (87x44.2x19.5 mm) (Coria and Currie, 2006)
(MMCH coll.) fourteen complete to fragmentary teeth (Neloadino, online 2016)
(MUCP uncatalogued) tooth (56x31.5x17 mm) (Coria and Currie, 2006)
Diagnosis-
(after Coria and Salgado, 1995) dorsoventrally wide main body of
maxilla with subparallel dorsal and ventral edges (unknown in Tyrannotitan); two pneumatic foramina on medial
surface of quadrate (unreported in Meraxes; unknown in Tyrannotitan); lobe-shaped obturator process.
Other diagnoses- Some
characters proposed by Coria and Salgado (1995) have since been
identified as diagnostic of all giganotosaurines and sometimes more
basal carcharodontosaurids- eave-like supraorbital lacrimal-postorbital
contact; symphyseal end of dentary dorsoventrally expanded, forming
ventral process; dorsally projected femoral head. The posterior
intercondylar groove in the proximal tibia is present in most
theropods. The tubercle-like insertion for triceps in the ventral
scapular edge is also present in Meraxes (but not Mapusaurus; unknown in Tyrannotitan).
Canale et al. (2015) noted the supposed scapula-coracoid suture is
actually a break in the coracoid, so that contra Coria and Salgado, the
proximal end of the scapula is not anteriorly projected above the
coracoid.
Comments- Carrano et al. (2012) report a femur length of 1.365 m for
the holotype and believe the skull to be less than 1.5 m long when reconstructed
correctly. Canale et al. (2022) used the almost complete skull of Meraxes to estimate a skull length of 1.62 m for Giganotosaurus' holotype. Coria and Currie (2006) state Mapusaurus specimen "MCF-PVPH-108.68 is a 1040 mm long tibia, which is 7% smaller than the tibia of Giganotosaurus" and "MCF-PVPH-108.202 is an 860 mm long fibula that is actually 2 cm longer than the fibula of the 12.2 m long Giganotosaurus."
The holotype was discovered in 1993 (Calvo, 1999) 12 km SW of El Chocon
and presented at the 1994 SVP by Coria and Salgado. Contrary to
the eventual description, a premaxilla, incomplete jugal, quadratojugal
and astragalus were reported. However, Eddy and Clarke (2011)
list the premaxilla as preserved as well as the ectopterygoid and
pterygoid. Unfortunately only the braincase (Coria and Currie,
2002) and endocast (Carabajal and Canale, 2010) have been described and
figured in detail, and as most of the presacral column was unprepared
in 1995, the numbers of cervicals and dorsals listed here are based on
photos of the holotype laid out on display at the MMCH. Note
that one tibia-fibula and the pedes in this display (as shown in Fig.
13 of Calvo, 1999) as well as the skull and forelimbs are artificial
however. Hendrickx et al. (2014) figure the quadrate in all views
(mislabeled Shaochilong
in the caption for Figure 11), Novas et al. (2013) figure the maxilla
and dentary, Eddy and Clarke (2011) figure the lacrimal and
ectopterygoid, and Calvo (1999) figures the scapulocoracoid (also in
Canale et al., 2015) and femur in medial view. Canale et al.
(2015) noted the scapulocoracoid is actually damaged, so that the
scapula-coracoid suture, low acromion and unexpanded distal end are not
real features. Calvo (1999) described a tooth (MUCPv-52) found in
1987 "on the coast of the Lake Ezquiel Ramos Mexia approximately 5 km.
south of El Chocon" as Giganotosaurus
"because it is laterally compressed and oval in cross-section", which
is typical of most Mesozoic theropods. He stated "the crown shows
close resemblance with that of the holotype", but never provided
details. Calvo (1989) initially described the dentary MUCPv-95
found in 1987 at Cerro Los Candelaros in an abstract. He noted it "has a very marked bulge on its lower edge, until now only faintly observed in Piatnitzkysaurus" and found that "The anterior end of the dentary is quadrangular in outline, a character not observed in other carnosaurs" (translated). He concluded that "The
preliminary study suggests that this remains would correspond to a new
species of Theropoda, whose jaw would be similar in size to that of Tyrannosaurus." Calvo (1999)
illustrated the dentary, which was later described in detail by Calvo
and Coria (2000) who found it was 8% larger than the type. Valais
and Apesteguia (2001) briefly described a tooth (MPCA coll.) in an
abstract from the then recently discovered La Buitrera locality of Rio
Negro "assignable to Giganotosaurus carolinii Coria and Salgado or a related form" "Given
its large size (estimated total length: 215 mm), lateral sinuosity,
denticular morphology, marginal wrinkles, and provenance"
(translated). Coria and Currie (2006) listed two uncatalogued
teeth, from the FPDM and MUCP, of unpublished provenance.
Finally, Neloadino (2016 online) photographed a display at the MMCH
labeled "several of the 60 teeth of the carnivorous Giganotosaurus carolinii about the approximate shape of the mouth" (translated).
The combination Carcharodontosaurus carolinii was first used by Figueiredo
(1998) (probably due to confusing Carcharodontosaurus and Giganotosaurus,
as it was said to be discovered in 1995 but found in Morocco), but was used
as an explicit new combination by Paul (2010). Paul's Carcharodontosaurus concept would encompass all of Carcharodontosaurinae as used here and has not been followed.
Initially Coria and Salgado (1994) only suggested "affinities with primitive tetanurines like Allosaurus",
while in its 1995 description they placed it closer to Avetheropoda
than Megalosauroidea without evidence of a quantified analysis, and
without justification for excluding it from Carnosauria. This was
apparently based on a 6 OTU 36 character quantified analysis (Coria,
1998), although the character list and data matrix were never
published. Sereno et al. (1996) were the first to recover Giganotosaurus in Carcharodontosauridae, where it has remained in virtually all later studies.
References-
Calvo, 1989. Un gigantesco theropodo del Miembro Candeleros
(Albiano-Cenomaniano) de la Formaci�n R�o Limay, Provincia del Neuqu�n,
Patagonia, Argentina. VII Jornadas Argentinas de Paleontolog�a de
Vertebrados. Ameghiniana. 26(3-4), 241.
Coria and Salgado, 1994. A giant theropod from the middle
Cretaceous of Patagonia, Argentina. Journal of Vertebrate Paleontology. 14(3),
22A.
Coria and Salgado, 1995. A new giant carnivorous dinosaur from the Cretaceous
of Patagonia. Nature. 377, 224-226.
Sereno, Dutheil, Iarochene, Larsson, Lyon, Magwene, Sidor, Varricchio and Wilson,
1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation.
Science. 272(5264), 986-991.
Coria, 1998. Relaciones filogeneticas de un gigantesco dinosaurio
teropodo del Cretacico de Patagonia. XI Jornadas Argentinas de
Paleontolog�a de Vertebrados (1995): Res�menes de trabajos presentados.
Acta Geological Lilloana. 18(1), 159-160.
Figueiredo, 1998. Os dinoss�urios carn�voros: A sua descri��o
e modo de vida. Centro Portugues de Geo-historia e Pre-historia. 4 pp.
Barrick and Showers, 1999. Thermophysiology and biology of Giganotosaurus: Comparison with Tyrannosaurus. Palaeontologia Electronica. 2.2.12A.
Calvo, 1999. Dinosaurs and other vertebrates of the Lake Ezequiel Ramos Mexia
area, Neuquen - Patagonia, Argentina. In Tomida, Rich and Vickers-Rich (eds.).
Proceedings of the Second Godwanan Dinosaur Symposium. 13-45.
Calvo and Coria, 2000. New specimen of Giganotosaurus carolinii (Coria & Salgado, 1995), supports
it as the largest theropod ever found. Gaia. 15, 117-122.
Blanco and Mazzetta, 2001. A new approach to evaluate the cursorial ability of the giant theropod Giganotosaurus carolinii. Acta Palaeontologica Polonica. 46(2), 193-202.
Valais and Apesteguia, 2001. Dientes asignables a Giganontosaurus (Carcharodontososauria,
Theropoda) provenientens de "La Buitera", Formacion Candeleros, provincia
de Rio Negro. Ameghiniana. 38(4), 6R-7R.
Coria and Currie, 2002. The braincase of Giganotosaurus carolinii (Dinosauria:
Theropoda) from the Upper Cretaceous of Argentina. Journal of Vertebrate Paleontology.
22(4), 802-811.
Mazzetta, Blanco and Cisilino, 2004. Modelizaci�n con elementos finitos de un diente referido al g�nero Giganotosaurus Coria y Salgado, 1995 (Theropoda: Carcharodontosauridae). Ameghiniana. 41(4), 619-626.
Mazzetta,
Christiansen and Farina, 2004. Giants and bizarres: Body size of some
southern South American Cretaceous dinosaurs. Historical Biology.
16(2-4), 71-83.
Coria and Currie, 2006. A new carcharodontosaurid (Dinosauria, Theropoda) from
the Upper Cretaceous of Argentina. Geodiversitas. 28(1), 71-118.
Carabajal and Canale, 2010. Cranial endocast of the carcharodontosaurid theropod
Giganotosaurus carolinii Coria & Salgado, 1995. Neues Jahrbuch f�r
Geologie und Pal�ontologie Abhandlungen. 258(2), 249-256.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Snively, Witmer, Ridgely, Wroe and Ryan, 2010. Impact and scythe-like jaw function in large Cretaceous theropods: Majungasaurus, Tyrannosaurus and Giganotosaurus compared. Journal of Vertebrate Paleontology. 28(3), 168A.
Eddy and Clarke, 2011. New information on the cranial anatomy of Acrocanthosaurus
atokensis and its implications for the phylogeny of Allosauroidea (Dinosauria:
Theropoda). PLoS ONE. 6(3), e17932.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Hendrickx, Araujo and Mateus, 2014. The nonavian theropod quadrate II:
Systematic usefulness, major trends and cladistic and phylogenetic
morphometrics analyses. PeerJ PrePrints. DOI:
10.7287/peerj.preprints.380v1
Canale, Novas and Pol, 2015. Osteology and phylogenetic relationships of Tyrannotitan
chubutensis Novas, de Valais, Vickers-Rich and Rich, 2005 (Theropoda: Carcharodontosauridae)
from the Lower Cretaceous of Patagonia, Argentina. Historical Biology. 27(1),
1-32.
Carabajal, Canale and Kundrat, 2015. Nueva informacion sobre la neuroanatomia
de Giganotosaurus carolinii usando tomografias computadas: Morfologia
del oido interno. XXIX Jornadas Argentinas de Paleontolog�a de Vertebrados,
resumenes. Ameghiniana. 52(4) suplemento, 63-64.
Neloadino, online 2016. https://upload.wikimedia.org/wikipedia/commons/2/22/Dientes_f%C3%B3siles_de_Giganotosaurus_en_el_museo_de_El_Choc%C3%B3n.JPG
Nieto and Paulina-Carabajal, 2020. 3D models related to the publication: Brief comment on the brain and inner ear of Giganotosaurus carolinii (Dinosauria: Theropoda) based on CT scans. MorphoMuseuM. DOI: 10.18563/journal.m3.108
Paulina-Carabajal and Nieto, 2020 (online 2019). Brief comment on the brain and inner ear of Giganotosaurus carolinii (Dinosauria: Theropoda) based on CT scans. Ameghiniana. 57, 58-62.
Canale, Apestegu�a, Gallina, Mitchell, Smith, Cullen, Shinya, Haluza,
Gianechini and Makovicky, 2022. New giant carnivorous dinosaur
reveals convergent evolutionary trends in theropod arm reduction.
Current Biology. 32(14), 3195-3202.
Allosauridae Marsh, 1878
Definition- (Allosaurus fragilis <- Sinraptor dongi,
Carcharodontosaurus saharicus) (Holtz et al., 2004)
Other definitions- (Allosaurus fragilis <- Sinraptor dongi)
(modified from Padian and Hutchinson, 1997)
(Allosaurus fragilis <- Sinraptor dongi, Monolophosaurus
jiangi, Cryolophosaurus ellioti, Carcharodontosaurus saharicus)
(modified from Sereno, 1998)
(Allosaurus fragilis <- Sinraptor dongi, Carcharodontosaurus
saharicus, Passer domesticus) (Brusatte and Sereno, 2008)
= Labrosauridae Marsh, 1882
= Antrodemidae Stromer, 1934
= Allosaurinae Marsh, 1878 sensu Paul, 1988
= Allosauridae sensu Sereno, 1998
Definition- (Allosaurus fragilis <- Sinraptor dongi, Monolophosaurus
jiangi, Cryolophosaurus ellioti, Carcharodontosaurus saharicus)
(modified)
= Allosauridae sensu Brusatte and Sereno, 2008
(Allosaurus fragilis <- Sinraptor dongi, Carcharodontosaurus
saharicus, Passer domesticus)
Comments- Brusatte and Sereno's (2008) definition differs from Holtz
et al.'s (2004) by including Passer as an external specifier. In this
case, I agree it's useful for cases like Paul (2002), Longrich (2001) and Coria
and Salgado (1995). I wonder if Tyrannosaurus might be a useful external
specifier as well, as tyrannosaurids and allosaurids have often been posited
as sister groups (Paul, 1988; Kurzanov, 1989; Molnar et al., 1990). However,
in all these cases, Carcharodontosaurus was seen as an intermediate between
Allosaurus and Tyrannosaurus, which would keep tyrannosaurids
from being allosaurids under Brusatte and Sereno's and Holtz et al.'s definitions.
undescribed possible allosaurid (Hand and Bakker, 2000)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Wyoming, US (Como Bluffs)
Material- ("larger than Allosaurus") (juvenile?) ulna,
radius, semilunate carpal, manus including manual ungual I
Diagnosis- (after Hand and Bakker, 2000) arm 25-30% shorter compared
to body than in Allosaurus; very robust; radius and ulna shorter than
manual ungual I (~185% longer in A. fragilis); manual ungual I long and
straight with large flexor tubercle.
Comments- This has only been mentioned in an abstract as a new allosaurid,
though the large size, robust and short arm, shortened forearm and enlarged
manual ungual I remind one of Torvosaurus. Note this is not "Wyomingraptor",
as it was discovered in 1999 while the nomen nudum was published in 1997.
References- Hand and Bakker, 2000. Implications of the functional morphology
of a new allosaurid forearm from the Como Bluffs. The Florida Symposium on Dinosaur
Bird Evolution. Publications in Paleontology No. 2, Graves Museum of Archaeology
and Natural History. 17.
unnamed allosaurid (Zinke, 1998)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Diagnosis- (after Zinke, 1998) differs from Allosaurus in lacking
downward-pointing blood grooves.
Material- (IPFUB GUI D 66) tooth (Rauhut, 2000)
(IPFUB GUI D 191-194) four teeth (~8.75 mm)
Comments- Chure (2000) doubted they could be referred to Allosauridae,
with the rather weak defense of "they are just more similar to Allosaurus
than other taxa", which would be in itself an acceptable reason to refer
them to this family.
References- Zinke, 1998. Small theropod teeth from the Upper Jurassic
coal mine of Guimarota (Portugal). Palaontologische Zeitschrift. 72, 179-189.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Rauhut, 2000. The dinosaur fauna from the Guimarota mine. In Martin and Krebs
(eds.). Guimarota, A Jurassic Ecosystem. Verlag Dr. Friedrich Pfeil. 75-82.
unnamed Allosauridae (Alcala, Royo-Torres, Cobos and Luque, 2009)
Late Tithonian-Middle Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (CPT-1326) tooth (16.5x8.5x5.1 mm)
(CPT-1437) tooth (~30.5x13x6.7 mm)
(CPT-1518) tooth (29.4x10.6x4.9 mm)
(CPT-1660) tooth (16x?x5.7 mm)
(CPT-2331) tooth (19.5x9.2x5.5 mm)
References- Alcala, Royo-Torres, Cobos and Luque, 2009. Updating dinosaur
record from Teruel (Aragon, Spain). Journal of Vertebrate Paleontology. 29(3),
53A.
Gasc�, Cobos, Royo-Torres, Mampel and Alcal�, 2012. Theropod teeth
diversity from the Villar del Arzobispo Formation (Tithonian-Berriasian) at
Riodeva (Teruel, Spain). Palaeobiodiversity and Palaeoenvironments. 92(2), 273-285.
unnamed possible allosaurid (Perez-Moreno, Sanz, Sudre and Sige, 1993)
Early Valanginian, Early Cretaceous
unnamed formation, Gard, France
Material- (MM-2) proximal manual phalanx II-2
(MM-8) fragmentary dorsal rib
(MM-9) fragmentary dorsal rib
(MM-11) manual ungual II
(MM-12) manual ungual III (~82 mm)
(MM-13) manual ungual III
(MM-14) manual phalanx III-2 (~50 mm)
(MM-15) manual phalanx III-3 (67 mm)
(MM-16) metacarpal III (10.2 mm)
(MM-17) manual ungual II (~114 mm)
(MM-18) manual phalanx II-1 (10.2 mm)
(MM-19) metacarpal I (82 mm)
(MM-20) humerus (~210 mm)
(MM-21) scapular fragment
Comments- Similar to Allosaurus in the medial concavity of metacarpal
I, in which it is unlike Acrocanthosaurus. Thus it is provisionally referred
to this family.
References- Perez-Moreno, Sanz, Sudre and Sige, 1993. A theropod dinosaur
from the Lower Cretaceous of Southern France. Revue de Paleobiologie. 7, 173-188.
undescribed Allosauridae (Gerke and Wings, 2014)
Kimmeridgian, Late Jurassic
Langenberg Quarry and/or Hannover, Germany
Material- (NLMH coll.) teeth
Reference- Gerke and Wings, 2014. Characters versus morphometrics: A
case study with isolated theropod teeth from the Late Jurassic of Lower Saxony,
Germany, reveals an astonishing diversity of theropod taxa. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 137.
undescribed allosaurid (Matsukawa and Obata, 1994)
Aptian-Albian, Early Cretaceous
Zouyun Formation, Mongolia
Comments- Matsukawa and Obata (1994) report through personal communication
with Mateer (1992) that an indeterminate allosaurid was discovered in the Aptian-Albian
Zouyun Formation of Mongolia.
Reference- Matsukawa and Obata, 1994. Cretaceous, a contribution to dinosaur
facies in Asia based on molluscan paleontology and stratigraphy. Cretaceous
Research. 15, 101-125.
unnamed possible allosaurid (Dong, 1997)
Barremian-Albian, Early Cretaceous
Xinminbao Group, Gansu, China
Material- (IVPP V.11122-3) tooth (24 mm)
Comments- Though referred to the Allosauridae by Dong (1997), Chure (2001)
found no support for this assignment and considered it Theropoda indet..
The tooth is moderately tall and recurved with both mesial and distal serrations.
The former may be confined to the apical half and seem subequal in size to the
distal ones.
References- Dong, 1997. On small theropods from Mazongshan Area, Gansu
Province, China. In Dong (ed.). Sino-Japanese Silk Road Dinosaur Expedition.
China Ocean Press, Beijing. 13-18.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
unnamed possible allosaurid (Park et al., 2000)
Hauterivian, Early Cretaceous
Hasandong Formation of the Shindong Group, South Korea
Material- (KPE 8004) tooth
(KPE 8005) tooth
References- Park, Yank, and Currie, 2000. Early Cretaceous dinosaur teeth
of Korea. In Lee (ed.). International Dinosaur Symposium for Kosong County in
Korea. Paleontological Society of Korea Special Publication. 4, 85-98.
Lee, 2003. Dinosaur bones and eggs in South Korea. Memoir of the Fukui Prefectural
Dinosaur Museum. 2, 113-121.
undescribed allosaurid (Serrano-Martinez, Ortega and Knoll, 2013)
Bathonian?, Middle Jurassic
Argiles de l'Irhazer of the Irhazer Group, Niger
Material- two teeth
Comments- Serrano-Martinez et al. (2013) noted these plot with allosaurids
when examined morphometrically.
Reference- Serrano-Martinez, Ortega and Knoll, 2013. Isolated theropod
teeth from the "Argiles de l'Irhazer" (Middle Jurassic) of Niger.
Journal of Vertebrate Paleontology. Program and Abstracts 2013, 210.
Allosaurus? ferox Marsh, 1896
= Antrodemus ferox (Marsh, 1896) Ostrom and McIntosh, 1966
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Quarry 14)
Holotype- (YPM 1893) partial skull (partial premaxillae, incomplete maxilla,
incomplete jugal, nasal fragment, lacrimal fragment, two cranial fragments),
dentaries (one partial, one incomplete), surangular fragment, dorsal central
fragment, dorsal rib fragments, (?)scapular fragment, incomplete pedal ungual
Comments- Allosaurus ferox (Marsh, 1896) is from the Tithonian
Brushy Basin Member (Quarry 14) of the Morrison Formation in Wyoming. It is
based on YPM 1893 (partial skull, partial dentaries, partial surangular, dorsal
central fragment, dorsal rib fragments, scapular fragment, incomplete pedal
ungual), which was found in 1882. Named in a footnote, Marsh only mentioned
that it has a maxillary fenestra unlike Ceratosaurus, which is now known
to be present in all Allosaurus specimens. Still, the few requirements
necessary for publication prior to 1930 means it's a valid name according to
the ICZN. Hay (1908) believed it to be a typo for Allosaurus fragilis.
It's been largely ignored thanks to its obscure origin and homonymy with Labrosaurus
ferox (USNM 2315 from Colorado), though generally has been considered a
junior synonym of fragilis (e.g. Molnar et al., 1990). Glut (1997) published
two photos of the specimen, and Chure (2000) has been the only author to describe
it. He states the few odd features (e.g. convex ventral maxillary margin) are
caused by incorrect restoration, though the antorbital fossa is better developed
than most specimens. Chure refers ferox to Allosaurus fragilis
as it supposedly has a highly angled ventral jugal margin unlike his A. jimmadseni, but his plate 102 shows this as restored and the angle actually varies within specimens referred to each species (see A. jimmadseni entry).
As no elements which differ between Allosaurus and Saurophaganax
are preserved, ferox is here viewed as Allosauridae indet..
References- Marsh, 1896. The dinosaurs of North America. United States
Geological Survey, 16th Annual Report, 1894-95. 55, 133-244.
Hay, 1908. On certain genera and species of carnivorous dinosaurs, with special
reference to Ceratosaurus nasicornis Marsh. Proceedings of the United
States National Museum. 35(1648), 351-366.
Ostrom and McIntosh, 1966. Marsh's Dinosaurs, the collection from Como Bluff.
Yale University Press, New Haven. 388 pp.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Osm�lska
and Dodson (eds.). The Dinosauria. University of California Press. 169-209.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076
pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Antrodemus Leidy, 1870
A. valens (Leidy, 1870) Leidy, 1870
= Poekilopleuron valens Leidy 1870
= Megalosaurus valens (Leidy, 1870) Nopsca, 1901
= Allosaurus valens (Leidy, 1870) Huene, 1932
Morrison Formation?, Middle Park, Grand County, Colorado, US
Late Jurassic?
Holotype- (USNM 218) partial ?sixth caudal centrum (~125 mm)
Diagnosis- Indeterminate at the level of Averostra but nearly identical
to the contemporaneous Allosaurus fragilis.
Comments- The holotype was discovered on July 26 or 27, 1869 (Burger, 2023) and described by Leidy
(1870) as a new species of Poekilopleuron (misspelled Poicilopleuron),
but stated if the trabeculae crossing the internal cavity is shown to be of
generic significance, "it might be named Antrodemus." This
single distinguishing feature is probably positional, as Poekilopleuron
only preserves more distal caudals, and Ceratosaurus shows trabeculae
in proximal caudals but a single central cavity in distal caudals. It was figured
as Antrodemus when in Leidy's (1873) followup paper. Nopcsa (1901) transferred
it to Megalosaurus without comment. Gilmore (1920) viewed it as synonymous
with Allosaurus based on a section of caudal six from USNM 8367, and
it is indeed basically identical. However, he does not also show they can be
distinguished from other genera. Matthew and Brown (1922) were the first to
note Allosaurus has the advantage of having a 'topotype' (USNM 4734)
from the same locality, while the locality of Antrodemus is inexact.
Future authors varied in their choice of Antrodemus vs. Allosaurus,
Huene (1932) using the impossible combination Allosaurus valens (impossible
as Antrodemus has priority). Madsen (1976) viewed Antrodemus as
"questionably" generically diagnostic and not specfically diagnostic,
though he distinguishes it from Ceratosaurus (narrow ventral edge, smaller
median groove, no paired camerae), he does not note any other taxa besides Allosaurus
that have similar caudals, and does not recognize other species of Allosaurus
besides A. fragilis. Thus neither Gilmore nor Madsen fully supported
their ideas of validity. Chure (2000) distinguishes Antrodemus from Ceratosaurus
and Torvosaurus (no pleurocoels), states caudals of Saurophaganax
"have been too badly damaged during preparation for comparison", and
notes definitely referred caudals of Marshosaurus and Stokesosaurus
are unknown. He believes the frequency of Allosaurus finds and similarity
with Antrodemus suggests these are synonymous genera, but believes it
can not be distinguished from A. fragilis or his new A. "jimmadseni".
Thus Antrodemus may turn out to be generically diagnostic but is not
specifically diagnostic in his view.
Comparing Antrodemus to Saurophaganax confirms the caudals don't
definitely differ, with the apparently taller centrum and missing lateral fossae
of the latter being potentially positional variation or due to poor preparation.
Marshosaurus-relative Condorraptor and Stokesosaurus-relative
Juratyrant show no obvious differences, with the minor differences (no
ventral groove in Condorraptor's proximal caudal, no obvious chevron
facet in Juratyrant's complete proximal caudal centrum) easily being
positional variation. Carcharodontosaurids differ in lacking ventral median
grooves, though Antrodemus cannot be distinguished from metriacanthosaurids
using published information. In conclusion it seems Antrodemus cannot
be classified more exactly than Averostra, though it can be distinguished
from many particular taxa. It's probable that if more taxa were sectioned or
CT scanned to determine their cross sectional shape and internal structure,
more could be determined. The near exact structural match, provenence and large
number of Allosaurus finds do suggest Antrodemus is the same taxon,
though this cannot yet be satisfactorily demonstrated. As the exact stratigraphy
is unknown, it's not even known if this would be referrable to the Brushy Basin
species or the Salt Wash one if Loewen's taxonomic ideas prove correct. As the
diagnostic USNM 4734 is likely to be designated the neotype of Allosaurus
fragilis shortly, Allosaurus fragilis will have no meaningful competition
to be the valid name for the Morrison allosaurid.
References- Leidy, 1870. Remarks on Poicilopleuron valens, Clidastes
intermedius, Leiodon proriger, Baptemys wyomingensis, and
Emys stevensonianus. Proceedings of the Academy of Natural Sciences of
Philadelphia. 22(1), 3-5.
Leidy, 1873. Contributions to the extinct vertebrate fauna of the Western territories.
Report of the United States Geological Survey of the Territories. 1-358.
Nopcsa, 1901. Synopsis und Abstammung der Dinosaurier. Foldtani kozlony. 31,
247-288.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. Bulletin of the United States National Museum. 110,
1-154.
Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus
from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History.
46(6), 367-385.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Madsen, 1976. Allosaurus fragilis: A revised osteology. Utah Geological
and Mineral Survey Bulletin. 109, 1-163.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Burger, 2023. Mystery in Middle Park: Relocating the site of Colorado's
first dinosaur discovery. Earth Sciences History. 42(1), 102-122.
Creosaurus Marsh, 1878
C. atrox Marsh, 1878
= Antrodemus atrox (Marsh, 1878) Gilmore, 1920
= Allosaurus atrox (Marsh, 1878) Paul, 1987
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Reed’s
Quarry 1)
Holotype- (YPM 1890) incomplete premaxilla, incomplete jugal, anterior tooth,
two lateral teeth, fragmentary anterior dorsal neural spine (?), partial ?twelfth
dorsal rib, two partial fused sacral vertebrae, third sacral rib, two proximal
to mid caudal centra (107, 100 mm), partial proximal caudal neural arch, incomplete
~twenty-third caudal neural arch, four mid to distal caudal central fragments,
incomplete ~forty-sixth caudal vertebra (78 mm), ?sternum (lost), incomplete
ilium (~710 mm), astragalus, distal tarsal III, ?metatarsal (277 mm; lost),
pedal phalanx II-1, pedal phalanges III-1, pedal ungual III, pedal phalanx IV-1
Comments- The type of Creosaurus atrox was discovered in 1878.
Note this discussion only concerns the holotype, and not the long-snouted morphology
called Allosaurus atrox or Creosaurus by e.g. Bakker and Paul
(discussed under "Short-snouted fragilis vs. long-snouted atrox?"
in the Allosaurus comments). Marsh originally diagnosed this taxon based
on trihedral (premaxillary) teeth with serrations, amphicoelous (posterior)
centra, elongate distal caudals, elongate metapodials (lost) and sharp unguals,
all now known to be standard for carnosaurs. Marsh reported at least one 277
mm long metatarsal, though it has not been commented on since and may be lost
prior to 1916 or misidentified. Note that the supposed posterior dorsal vertebra
(actually a proximal caudal) identified by Marsh (1884) as Creosaurus
is actually YPM 1932. Chure (2000) notes the hyoid and manual ungual listed
by Lull (1916 pers. comm. to Gilmore, 1920) are part of the Labrosaurus lucaris
type, while the supposed sternal cannot be reconciled with existing specimens.
As early as Williston (1878), workers supposed Creosaurus might be synonymous
with Allosaurus, with supposed pectoral and forelimb differences mentioned
by Williston (1901) and Hay (1908) being due to Marsh's 'Allosaurus'
forelimb being a composite with Ceratosaurus and Megalosaurus
elements. Osborn (1903) described skulls AMNH 600 and 666 as Creosaurus,
which probably gave birth to the concept of long-snouted Creosaurus adopted
by Bakker and Paul decades later. There was no justification for referring either
skull to Creosaurus, and Osborn referred both to Allosaurus a
few years later (Osborn, 1906) and from then on. Gilmore (1920) agreed Creosaurus
was a junior synonym of Allosaurus due to a lack of published differences,
though he declined to discuss specific synonymization pending further study.
Paul and Carpenter (2010) merely stated it was a nomen dubium without evidence.
There had been no examination of the Creosaurus holotype after this until
Chure's thesis, where it is redescribed and partially illustrated. Note an anterior
dorsal neural spine fragment is listed as part of the type, but not described.
Chure referred YPM 1890 to Allosaurus sp., stating it "does not
show any feature to differentiate it from Allosaurus fragilis or A.
jimmadseni", despite the fact the jugal shows the ventral deflection
he believes characterizes A. fragilis and the ilium has a vertical ridge
that he states "is not present on any A. fragilis specimen"
but is on the jimmadseni holotype. Chure also notes that the
second and third sacrals of atrox are V-shaped ventrally in section,
unlike those of other Allosaurus specimens. The fourth and fifth sacrals
of jimmadseni's type have this type of venter, but the fourth sacral
centra of all Allosaurus specimens have a large lateral foramen, unlike
these sacrals. So the sacrum is different from other Allosaurus specimens,
no matter which vertebrae are represented. YPM 1890 thus seems to be an intermediate
specimen that helps decrease the difference between fragilis and jimmadseni.
Finally, Chure states pedal ungual III differs from A. fragilis in having
a proximally directed traingular tuber on the proximodorsal surface, though
this seems to be easily explainable by individual variation. In relation to
the larger but coeval Saurophaganax, Creosaurus doesn't preserve
any elements which are said to differ between it and Allosaurus. It is
thus Allosauridae indet., especially as the incomplete sacral and proximal/mid
caudal fusion could indicate immaturity.
References- Marsh, 1878. Notice of new dinosaurian reptiles. American
Journal of Science and Arts. 15, 241-244.
Williston, 1878. American Jurassic dinosaurs. Transactions of the Kansas Academy
of Science. 6, 42-46.
Marsh, 1884. Principal characters of American Jurassic dinosaurs. Part VIII.
The order Theropoda. The American Journal of Science, series 3. 27, 329-340.
Williston, 1901. The dinosaurian genus Creosaurus, Marsh. American Journal
of Science, series 4. 11, 111-114.
Osborn, 1903. The skull of Creosaurus. Bulletin of the American Museum
of Natural History. 19(31), 697-701.
Osborn, 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur (Second
communication). Bulletin of the American Museum of Natural History. 22(16),
281-296.
Hay, 1908. On certain genera and species of carnivorous dinosaurs, with special
reference to Ceratosaurus nasicornis Marsh. Proceedings of the United
States National Museum. 35(1648), 351-366.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. Bulletin of the United States National Museum. 110,
1-154.
Paul, 1987. The science and art of restoring the life appearance of dinosaurs
and their relatives: A rigorous how-to guide. In Czerkas and Olson (eds.). Dinosaurs
Past and Present. 2, 4-49.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Paul and Carpenter, 2010. Allosaurus Marsh, 1877 (Dinosauria, Theropoda):
Proposed conservation of usage by designation of a neotype for its type species
Allosaurus fragilis Marsh, 1877. Bulletin of Zoological Nomenclature.
67(1), 53-56.
Epanterias Cope, 1878
E. amplexus Cope, 1878
= Allosaurus amplexus (Cope, 1878) Paul, 1988
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US (Cope's Nipple)
Holotype- (AMNH 5767) axis, sixth or seventh cervical centrum (115 mm),
first dorsal neural arch, coracoid, distal metatarsal IV
Referred- ?(CPS 99) proximal caudal vertebra, distal caudal vertebra
(Bakker, 1990)
Diagnosis- Indeterminate relative to Allosaurus and Saurophaganax.
Comments- Epanterias amplexus (Cope, 1878) is from the Tithonian
Brushy Basin Member (Cope's Nipple) of the Morrison Formation in Colorado. The
holotype is AMNH 5767 (an axis, mid cervical centrum, first dorsal neural arch,
coracoid and distal metatarsal IV) and was found in 1877. Only the centrum and
neural arch are referenced by Cope, who believed the taxon to be a camarasaurid.
The cited vertebral characters are typical of Allosaurus- strongly opisthocoelous
cervical with prominent anteriorly placed parapophyses; unelevated, low, wide
and anteroposteriorly short dorsal neural arch; no hyposphene before mid dorsals;
transversely elongate anterior dorsal neural spine which has three apices; thin
anterior dorsal infradiapophyseal laminae; three infradiapophyseal fossae of
which the infraprezygapophyseal fossa is vertical; deep postspinal fossa; postzygapophyses
which overhang the centrum. However, the claim the anterior dorsal centrum lacks
pleurocoels is untrue (Chure, 2000). Osborn and Mook (1921) were the first to
recognize that Epanterias was a theropod which "at present cannot
be separated from Allosaurus" and illustrate some of the remains.
They also provisionally referred ribs initially allocated to Camarasaurus
and a femur originally referred to Laelaps trihedrodon by Cope, though
Chure says these cannot be located in the AMNH collections. Their identity remains
uncertain. Paul (1988) referred amplexus questionably to Allosaurus
and thought Saurophaganax was a junior synonym, based on their shared
large size and high stratigraphical position, though he did not cite morphological
characters supporting this. Bakker (1990) followed this theory and separated
Epanterias as a different genus, referring two caudal vertebrae to it.
This was presumably based on size and stratigraphy, but as no caudals are preserved
in the type they cannot be compared. Contradicting Paul's and Bakker's hypothesis,
modern stratigraphical work has placed the average-sized Allosaurus ferox
type between Saurophaganax and Epanterias, and the more recently
discovered average-sized UMNH VP 5918-5926 is between them as well. Not only
that, the recently discovered large allosaur NMMNH P-26083 is stratigraphically
very close to the Cleveland-Lloyd Quarry which is full of average-sized specimens.
While Paul is correct that large allosaurs are too rare to be adults of average-sized
allosaurs, Epanterias and NMMNH P-26083 could simply be large individuals
or representatives of larger-sized populations. Neither were discovered with
average-sized allosaurs in the same quarry, and Bybee et al. (2006) state "Individuals
of Allosaurus did not all grow alike" and could not "directly
test whether Allosaurus had indeterminate growth." Paul and Carpenter
(2010) oddly stated Epanterias "is probably a different taxon from
YPM 1930" though also "a nomen dubium because holotype AMNH 5767 ...
is insufficiently diagnostic.", with neither assertion backed up with evidence.
Chure has been the only author to describe Epanterias in detail and compare
it to Saurophaganax and Allosaurus. Compared to the former,
it has supposedly differently oriented cervical parapophyses and no dorsal paraspinal
lamina. However, I see no difference in parapophyseal orientation (both ~40
degrees anterodorsally) and Chure admits that the Epanterias vertebra
is more anterior than known Saurophaganax dorsals with paraspinal laminae,
so it may not have developed until more posterior dorsals. Differences with
Allosaurus fragilis that Chure notes are a less laterally compressed
axial centrum, less rectangular distal outline of metatarsal IV and better developed
lateral condyle in that element. Yet the axial centrum seems equally compressed
in USNM 4734, and the metatarsal differences were made with comparison to Madsen's
composite illustrations. Differences with Allosaurus were considered
minor by Chure in any case, and he synonymized Epanterias with Allosaurus
fragilis. I agree they are minor if they exist, but also believe it cannot
be distinguished from Saurophaganax as noted above. Thus Epanterias
amplexus becomes Allosauridae indet. until further distinguishing characters
are found.
References- Cope, 1878. A new opisthocoelous dinosaur. American Naturalist.
12(6), 406.
Osborn and Mook, 1921. Camarasaurus, Amphicoelias, and other sauropods
of Cope. Memoirs of the American Museum of Natural History. New Series 3(3),
247-387.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Bakker, 1990. A new Latest Jurassic vertebrate fauna from the highest levels
of the Morrison Formation at Como Bluff, Wyoming, with comments on Morrison
biochronology. Part 1: Biochronology. Hunteria. 2(6), 1-3.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Bybee, Lee and Lamm, 2006. Sizing the Jurassic theropod dinosaur Allosaurus:
Assessing growth strategy and evolution of ontogenetic scaling of limbs. Journal
of Morphology. 267(3), 347-59.
Paul and Carpenter, 2010. Allosaurus Marsh, 1877 (Dinosauria, Theropoda):
Proposed conservation of usage by designation of a neotype for its type species
Allosaurus fragilis Marsh, 1877. Bulletin of Zoological Nomenclature.
67(1), 53-56.
Labrosaurus Marsh, 1879
L. lucaris (Marsh, 1878) Marsh, 1879
= Allosaurus lucaris Marsh, 1978
= Antrodemus lucaris (Marsh, 1878) Hay, 1902
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Reed’s
Quarry 3)
Holotype- (YPM 1931; holotype of Allosaurus lucaris) incomplete
eighth cervical vertebra, ninth cervical centrum, tenth cervical centrum, second
dorsal centrum, fused partial fourth and fifth dorsal vertebrae, neural spine
and zygapophyseal fragments, proximal scapulae, incomplete coracoids, humeri
(~322 mm), partial ulna, incomplete metacarpal II, phalanx II-1, proximal phalanx
II-2
....(YPM 46147) manual ungual
....(YPM coll.) surangular, prearticular, articular, hyoid, three teeth, first
dorsal centrum, third dorsal centrum, partial sixth dorsal vertebra, seventh
dorsal centrum, eighth dorsal centrum, two partial posterior dorsal vertebrae,
posterior dorsal centrum, incomplete posterior dorsal rib, five partial proximal
caudal vertebrae, three distal caudal centra, twelve distal caudal central fragments,
partial neural arch, transverse process, zygopophyseal fragments
Comments- Labrosaurus lucaris is from the Tithonian Brushy Basin
Member (Reed's Quarry 3) of the Morrison Formation in Wyoming. The holotype
YPM 1931 was discovered in 1878. It consists of a partial mandible, hyoid, partial
posterior cervical, dorsal and caudal vertebrae, many vertebral fragments, a
humerus and partial forelimb. Allosaurus lucaris was originally described
and diagnosed based solely on an anterior dorsal vertebra, which was diagnosed
by its opisthocoelous centrum with pleurocoels, ventral keel and anteriorly
placed parapophyses that are the usual characters of carnosaurs. Marsh (1879)
later made lucaris the type of his new genus Labrosaurus, in his
new family Allosauridae. He diagnosed it by short and strongly opisthocoelous
cervicals, moderately opisthocoelous (anterior) dorsals, both with pleurocoels
connected to large internal cavities, tall and transversely expanded neural
spines (in posterior cervicals and anterior dorsals), anterior presacrals without
hyposphenes, small forelimbs, curved humerus, and large deltopectoral ("radial")
crest, all now known in Allosaurus. Gilmore (1920) illustrated the humerus
in his plate 6 as Antrodemus valens, though otherwise it has only been
illustrated by Chure (2000). Synonymized with 'Antrodemus valens' (Allosaurus
fragilis) in Steel's (1970) review, it has been assumed to be a junior synonym
since. Chure also claims it shows no unique characters and refers it to A.
fragilis, though he does not distinguish it from A. jimmadseni
or even Saurophaganax. Paul and Carpenter (2010) oddly state it "probably
represents a different species from YPM 1930 [the Allosaurus holotype],
and may or may not belong to the same genus. It is not sufficiently complete
to characterise a species." without defending any of these assertions.
Review of the published information indicates pleurocoels are reported in the
fifth dorsal ("pectoral 5" of Chure) as in Saurophaganax, yet
the proximal outline of manual phalanx II-2 resembles Allosaurus more
in being rectangular instead of oval. Either of these could easily be explained
by individual variation, and Labrosaurus is here assigned to Allosauridae
indet. pending further study.
References- Marsh, 1878. Notice of new dinosaurian reptiles. American
Journal of Science and Arts. 15, 241-244.
Marsh, 1879, Principal characters of American Jurassic dinosaurs. Part 1. American
Journal of Science and Arts. 16, 411-416.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 1-868.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. Bulletin of the United States National Museum. 110,
1-154.
Steel, 1970. Saurischia. Handbuch der Palaoherpetologie. 14, 1-87.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Paul and Carpenter, 2010. Allosaurus Marsh, 1877 (Dinosauria, Theropoda):
Proposed conservation of usage by designation of a neotype for its type species
Allosaurus fragilis Marsh, 1877. Bulletin of Zoological Nomenclature.
67(1), 53-56.
Saurophaganax Chure, 1995
= "Saurophagus" Stovall vide Ray, 1941
S. maximus Chure, 1995
= "Saurophagus maximus" Stovall vide Ray, 1941 (preoccupied Swainson,
1831)
= Allosaurus maximus (Chure, 1995) Smith, 1998
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Oklahoma, US
Holotype- (OMNH 1123) fifth dorsal neural arch
Paratypes- (OMNH 780) manual ungual I
(OMNH 1102) mid chevron
(OMNH 1135) atlas
(OMNH 1142) partial quadrate
(OMNH 1152) lateral tooth
(OMNH 1153) lateral tooth
(OMNH 1190) posterior dorsal centrum
(OMNH 1191) metatarsal III
(OMNH 1307) metatarsal II
(OMNH 1338) partial ilium, pedal ungual IV
(OMNH 1370) tibia (955 mm)
(OMNH 1396) metatarsal IV
(OMNH 1425) distal pubis
(OMNH 1438) mid chevron
(OMNH 1444) mid cervical vertebra
(OMNH 1680) lateral tooth
(OMNH 1681) distal metatarsal I
(OMNH 1685) mid chevron
(OMNH 1708) femur (1059 mm)
(OMNH 1737) proximal ischium
(OMNH 1771) postorbital
(OMNH 1935) humerus (545 mm)
(OMNH 2145) partial quadrate
(OMNH 4666; lectotype of "Saurophagus maximus") tibia
Referred- (OMNH 1104) mid chevron (Chure, 2000)
(OMNH 1122) proximal caudal neural arch (Chure, 2000)
(OMNH 1124) metacarpal III (Chure, 2000)
(OMNH 1126) pedal phalanx III-3 (Chure, 2000)
(OMNH 1127) manual phalanx III-1 (Chure, 2000)
(OMNH 1128) incomplete manual ungual III (Chure, 2000)
(OMNH 1180) partial mid chevron (Chure, 2000)
(OMNH 1189) dorsal centrum (Chure, 2000)
(OMNH 1192) metatarsal III (Chure, 2000)
(OMNH 1193) metatarsal IV (Chure, 2000)
(OMNH 1306) metatarsal IV (Chure, 2000)
(OMNH 1364) radius (Chure, 2000)
(OMNH 1415) radius (Chure, 2000)
(OMNH 1424) proximal ischium (Chure, 2000)
(OMNH 1426) incomplete fibula (Chure, 2000)
(OMNH 1433) two anterior sacral centra, sacral rib (Chure, 2000)
(OMNH 1434) incomplete ulna (Chure, 2000)
(OMNH 1439) mid chevron (Chure, 2000)
(OMNH 1446) partial posterior cervical centrum (Chure, 2000)
(OMNH 1447) dorsal centrum (Chure, 2000)
(OMNH 1450) anterior dorsal centrum (Chure, 2000)
(OMNH 1461) metatarsal II (Chure, 2000)
(OMNH 1684) mid chevron (Chure, 2000)
(OMNH 1693) distal humerus (Chure, 2000)
(OMNH 1694) incomplete fibula (Chure, 2000)
(OMNH 1695) incomplete fibula (Chure, 2000)
(OMNH 1703) proximal ischium (Chure, 2000)
(OMNH 1707) proximal pubis (Chure, 2000)
(OMNH 1904) proximal caudal vertebra (Chure, 2000)
(OMNH 1906) anterior dorsal centrum (Chure, 2000)
(OMNH 1911) pedal phalanx IV-1 (Chure, 2000)
(OMNH 1912) pedal ungual IV (Chure, 2000)
(OMNH 1913) pedal ungual IV (Chure, 2000)
(OMNH 1914) pedal ungua IV (Chure, 2000)
(OMNH 1915) pedal ungual I (Chure, 2000)
(OMNH 1916) pedal phalanx III-1 (Chure, 2000)
(OMNH 1918) pedal phalanx II-1 (Chure, 2000)
(OMNH 1919) pedal phalanx III-2 (Chure, 2000)
(OMNH 1920) manual phalanx II-2 (Chure, 2000)
(OMNH 1921) manual phalanx II-1 (Chure, 2000)
(OMNH 1924) metatarsal III (Chure, 2000)
(OMNH 1925) pedal phalanx III-3 (Chure, 2000)
(OMNH 1927) mid caudal vertebra (Chure, 2000)
(OMNH 1928) mid caudal vertebra, metacarpal I (Chure, 2000)
(OMNH 1929) metacarpal II (Chure, 2000)
(OMNH 1936) metatarsal IV (Chure, 2000)
(OMNH 1947) fifth sacral vertebra (Chure, 2000)
(OMNH 2114) femur (Chure, 2000)
(OMNH 2146) mid cervical vertebra (Chure, 2000)
(OMNH 2147) mid cervical vertebra (Chure, 2000)
(OMNH 2149) distal tibia (Chure, 2000)
(OMNH 2154) incomplete scapula (Chure, 2000)
(OMNH 4016) partial scapula (Chure, 2000)
(OMNH 10357) proximal caudal vertebra (Chure, 2000)
(OMNH 10373) pedal phalanx III-1 (Chure, 2000)
(OMNH 10375) pedal phalanx IV-2 (Chure, 2000)
(OMNH 10376) pedal phalanx I-1 (Chure, 2000)
(OMNH 10377) pedal phalanx I-1 (Chure, 2000)
(OMNH 10381) femur (Chure, 2000)
(OMNH 10732) pedal phalanx III-1 (Chure, 2000)
(OMNH 52384) pedal phalanx II-2 (Chure, 2000)
(OMNH 52385) pedal phalanx II-1 (Chure, 2000)
(OMNH 52386) pedal phalanx III-2 (Chure, 2000)
(OMNH 52387) pedal phalanx III-2 (Chure, 2000)
(OMNH 52388) pedal phalanx III-3 (Chure, 2000)
(OMNH 52389) pedal phalanx IV-1 (Chure, 2000)
(OMNH 52390) pedal phalanx IV-2 (Chure, 2000)
(OMNH 52391) pedal phalanx IV-3 (Chure, 2000)
(OMNH 52392) pedal phalanx IV-3 (Chure, 2000)
(OMNH 52393) pedal ungual II (Chure, 2000)
(OMNH 52394) pedal ungual II (Chure, 2000)
(OMNH 52395) pedal ungual III (Chure, 2000)
(OMNH coll.) posterior dorsal centra, metatarsal II, pedal phalanx III-1 (Chure,
2000)
Diagnosis- (after Chure, 1995) no atlantal prezygapophyses (in OMNH 1135);
no medial projection from atlantal arch to roof over neural canal (in OMNH 1135);
horizontal lamina along base of each side of mid dorsal neural spines arising
from spine base anteriorly, free posteriorly (in OMNH 1123); ventral part of
chevrons greatly expanded anteriorly (in up to six specimens); femur bowed laterally
(in up to three specimens).
(after Chure, 2000) postorbital lacks rugosity (in OMNH 1771); square atlantal
intercentrum (in OMNH 1135); mid cervicals with nearly vertical postzygapophyses;
mid dorsals have well developed infraprezygapophyseal lamina (in OMNH 2146);
mid dorsals with vertically oriented infrapostzygapophyseal lamina (in OMNH
1123); pleurocoels present through fifth dorsal centrum (in OMNH 1123);
radius with marked elliptical muscle attachment near proximoventral margin of
lateral surface (in up to two specimens); spiral muscle scar on lateral surface
of radius shaft (in up to two specimens); proximal end of manual phalanx II-2
oval instead of rectangular (in OMNH 1921); weaker astragalar buttress on anterodistal
tibia (in both OMNH 1370 and 2149); medial malleolus projects twice as far medially
than Allosaurus (in up to two specimens); distal end of metatarsal IV
rectangular instead of triangular (in up to three specimens).
Other diagnoses- Chure (2000) stated that the cnemial crest is deflected
laterally more than in Allosaurus, the lateral surface of the lateral
tibial condyle is concave (instead of convex), the lateral tibial condyle is
more posteriorly placed than in Allosaurus, the fibular crest is more
set off from shaft, and the medioventral corner of pedal phalanx III-1's proximal
edge being angled inward were all unlike Allosaurus, but that genus shows
these conditions too (UUVP coll. from Madsen, 1976). Metatarsal IV is indeed
less distally divergent than some A. fragilis (e.g. AMNH 324, DINO 11541),
but not other fragilis specimens (e.g. MOR 693).
Comments- At least four individuals are represented by the OMNH material.
Chure (1995) originally mentioned anterior cervicals, caudal centra and a tridactyl
manus, which would be paratype material if their specimen numbers could be determined.
Though generally accepted as a species distinct from A. fragilis, Saurophaganax
is represented by multiple individuals with uncertain referral of particular
elements to each one, and reportedly distinctive characters are not stated to
be present in multiple specimens except for distal chevron expansion and astragalar
buttress strength. Indeed, nine of the characters can only be evaluated for
one specimen. Thus the apparent distinctiveness of this taxon may be due to
individuals each having a few unique characters, as is true of most well described
Morrison allosaurs. Resolution will involve more detailed information on each
specimen and more description of the variation in Morrison allosaurids. Smith
(2008) found no morphometric differences between maximus and fragilis
in at least the humerus and femur.
References- Ray, 1941. Big for his day. Natural History. 48, 36-39.
Camp, Welles and Green, 1953. Bibliography of fossil vertebrates 1944-1948.
Geological Society of America Memoir. 57, 465 pp.
Chure, 1995. A reassessment of the gigantic theropod Saurophagus maximus
from the Morrison Formation (Upper Jurassic) of Oklahoma, USA. Sixth Symposium
on Mesozoic Terrestrial Ecosystems and Biota. 103-106.
Smith, 1998. A morphometric analysis of Allosaurus. Journal of Vertebrate
Paleontology. 18(1), 126-142.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Allosaurus Marsh, 1877
pr= "Madsenius" Lambert, 1990
pr= "Wyomingraptor" Anonymous, 1997
Diagnosis- (modified from Chure, 2000) fenestrate dorsal wall of maxillary
antrum (unknown in Saurophaganax); spindle-shaped foramen on lateral
surface of sacral centrum 4 (unknown in Saurophaganax); obturator process
with long anteriorly directed lamina that extends to level of puboischiadic
contact (unknown in Saurophaganax).
(after Carrano et al., 2012) tall, transversely compressed lacrimal horn (unknown
in Saurophaganax); reduced external mandibular fenestra (unknown in Saurophaganax);
strongly downturned paraoccipital processes that terminate well ventral to basal
tubera (unknown in Saurophaganax); antarticular bone in mandible (unknown
in Saurophaganax); distal expansion of ischium suboval in lateral view
(unknown in Saurophaganax).
Not Allosaurus- The AMNH online catalogue lists AMNH 3838 as Allosaurus?
sp. based on several teeth from the Belly River Group of Alberta, Canada, but
these are near certainly tyrannosaurid based on locaility. Similarly, the UCMP
online catalogue lists UCMP 136555 and 136556 as Allosaurus based on
two teeth from the Judith River Group of Montana, which are also presumably
tyrannosaurid.
References- Marsh, 1877. Notice of new dinosaurian reptiles from the
Jurassic formation. American Journal of Science and Arts. 14, 514-516.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
A. fragilis Marsh, 1877
?= Camptonotus amplus Marsh, 1879
= Labrosaurus ferox Marsh, 1884
?= Camptosaurus amplus (Marsh, 1879) Marsh, 1885
= Labrosaurus fragilis (Marsh, 1877) Nopsca, 1901
= Antrodemus fragilis (Marsh, 1877) Lapparent and Zbyszewski, 1957
= Allosaurus "whitei" Pickering, 1995
= Allosaurus "jimmadseni" Chure, 2000a vide Glut, 2003
= Allosaurus "carnegeii" Levin, 2003
pr= "Madsenius trux" Bakker vide Williams, 2004 online
?= Allosaurus europaeus Mateus, Walen and Antunes, 2006
= Allosaurus lucasi Dalman, 2014
?= Allosaurus amplus (Marsh, 1879) Galton, Carpenter and Dalman, 2015
= Allosaurus jimmadseni Chure and Loewen, 2020
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US (Dry Mesa Quarry,
Marsh-Felch Quarry 1, Marsh-Felch Quarry 2, Mygatt-Moore Quarry, Wolf Creek Quarry)
Holotype- (YPM 1930) incomplete lateral tooth (~55 mm), incomplete cervical
or anterior dorsal centrum, incomplete ~fourth dorsal centrum (85 mm), incomplete
fifth sacral centrum (105 mm), two dorsal rib fragments, humeral fragment, pedal
phalanx III-1 (~109 mm)
Referred- (AMNH 5780) five teeth (Chure, 2001)
(BYU 725/?) jugal, humerus (Smith et al., 1999)
(BYU 725/5097) humerus (Smith et al., 1999)
(BYU 725/5098) humerus (Smith et al., 1999)
(BYU 725/9249) postorbital (Smith et al., 1999)
(BYU 725/5126) maxilla (Eddy and Clarke, 2011)
(BYU 725/10296) humerus (Smith et al., 1999)
(BYU 725/10602) dentary (Smith et al., 1999)
(BYU 725/13259) mid caudal vertebra (Malafaia et al., 2017)
(BYU 725/15599) axis (Smith et al., 1999)
(BYU 725/17125) dentary (Therrien et al., 2005)
(BYU 725/17879) braincase (Eddy and Clarke, 2011)
(BYU 4861) (Chure and Loewen, 2020)
(BYU 4878; = BYU 4878 of Smith et al., 1999) humerus (Smith et al., 1999)
(BYU 4891; = BYU 4981 of Chure, 2000a) pubes (Chure, 2000a)
(BYU 5099) humerus (Taylor, 1992)
(BYU 5122) material including jugal (Chure and Loewen, 2020)
(BYU 5125) lacrimal (Britt, 1991)
(BYU 5164) (Carrano et al., 2012)
(BYU 5268) (Carrano et al., 2012)
(BYU 5292) (Chure and Loewen, 2020)
(BYU 5583; = Mes 5583; typo for UUVP 5583?) (Carrano et al., 2012)
(BYU 11936; = Mes 11936) (Carrano et al., 2012)
(BYU 13621; = Mes 13621) (Carrano et al., 2012)
(BYU 13807) material including jugal (Chure and Loewen, 2020)
(BYU 16942; = Mes 16942) (Carrano et al., 2012)
(BYU 17106; = Mes 17106) (Carrano et al., 2012)
(BYU 17281; = Mes 17281) (Carrano et al., 2012)
(BYU coll.) nearly 200 elements (Britt, 1991)
(BYU coll.) (at least 13 individuals) 988 elements including femora (~120-960
mm) (Dangerfield, Britt and Scheetz, 2006)
....(BYU coll.) skull, caudal vertebrae, scapulocoracoid and hindlimbs including
femur (960 mm)
(DMNH 2734/HEC 647, 648) eggshells (Hirsch, 1994)
(DMNH coll.) (Lederer and Small, 1999)
(MWC 732.002.OO1/HEC 575) eggshells (Hirsch, 1994)
(MWC coll.) ~222 teeth (5-23 mm), skeletal elements (Kirkland and Armstrong,
1992)
(MWC coll.) seven teeth (Kane, 2020)
(UCM 54471/HEC 268) eggshell (Hirsch, 1994)
(USNM 4734; proposed neotype) (7.4 m, 1.01 tons) incomplete skull (682
mm), partial mandible, atlas (28 mm), axis (85 mm), third cervical
vertebra (105 mm), third cervical rib (268 mm), fourth cervical
vertebra (102 mm), fourth cervical rib, fifth cervical vertebra (109
mm), fifth cervical rib, sixth cervical vertebra (111 mm), sixth
cervical rib, seventh cervical vertebra (115 mm), eighth cervical
vertebra (115 mm), eighth cervical rib, ninth cervical vertebra (123
mm), ninth cervical rib, (dorsal series ~1107 mm) first dorsal vertebra
(88 mm), second dorsal vertebra (77 mm), third dorsal vertebra (74 mm),
fourth dorsal centrum (72 mm), fifth dorsal vertebra (74 mm), sixth
dorsal vertebra (81 mm), seventh dorsal vertebra (~81 mm), ninth dorsal
vertebra (85 mm), tenth dorsal vertebra (94 mm), eleventh dorsal
vertebra (93 mm), twelfth dorsal vertebra (99 mm), thirteenth dorsal
vertebra (106 mm), fourteen dorsal ribs, most gastralia, (sacrum 536
mm) first sacral vertebra (116 mm), second sacral vertebra (104 mm),
third sacral vertebra (104 mm), fourth sacral vertebra (108 mm), fifth
sacral vertebra (104 mm), thirty-three caudal vertebrae, six chevrons,
scapulae, coracoids, furcula, humeri (310 mm), radii (222 mm), ulnae
(263 mm), radiales, intermedium, distal carpals I, distal carpal II,
metacarpal I (73 mm), phalanges I-1 (136,138 mm), manual unguals I
(118, 120 mm), metacarpals II (125, 122 mm), phalanges II-1 (94, 94
mm), phalanx II-2 (102 mm), manual ungual II (95 mm), metacarpals III
(105, 97 mm), phalanges III-1 (50, 42 mm), phalanges III-2 (41, 43 mm),
phalanges III-3 (52, 55 mm), manual unguals III (61, 59 mm), ilia (672,
720 mm), pubes (680 mm), ischia (575 mm), femora (770 mm), tibiae (690
mm), fibulae (623 mm), astragali (132 mm wide), calcanea, distal tarsal
III, distal tarsal IV, metatarsal I (85 mm), phalanx I-1 (70 mm), pedal
ungual I (~70 mm), metatarsal II (270 mm), phalanx II-1 (120 mm),
phalanx II-2 (80 mm), metatarsal III (327 mm), phalanx III-1 (110 mm),
phalanx III-2 (~90 mm), phalanx III-3 (66 mm), metatarsal IV (275 mm),
phalanx IV-1 (75 mm), phalanx IV-2 (50 mm), phalanx IV-3 (30 mm),
phalanx IV-4 (29 mm) (Marsh, 1884; described by Gilmore, 1915, 1920)
?...(USNM 2315; holotype of Labrosaurus ferox) dentary (412 mm) (Marsh,
1884)
(USNM 7336) astragalus (208 mm wide, 172 mm high) (Gilmore, 1920)
(USNM 8335) maxilla, teeth, dentary (Gilmore, 1920)
(USNM 8423) maxillae, five dorsal centra, (sacrum 540 mm), first sacral vertebra
(~90 mm), second sacral vertebra (99 mm), third sacral vertebra (104 mm), fourth
sacral vertebra (120 mm), fifth sacral vertebra (114 mm), three caudal centra,
humerus, manual bones, partial ilia, broken pubes, broken ischia, femora (805
mm), metatarsal II (320 mm), phalanx II-1 (122 mm), metatarsal III (353 mm),
phalanx III-1 (116 mm), phalanx III-2 (94 mm), phalanx III-3 (74 mm), metatarsal
IV (324 mm), phalanx IV-2 (72 mm), pedal ungual IV (~75 mm) (Gilmore, 1920)
(YPM 57589; holotype of Allosaurus lucasi) (large) incomplete premaxilla,
premaxillary tooth, partial maxilla, quadratojugal, partial ?quadrate, braincase,
dentary fragment, several lateral teeth, incomplete ~nineteenth caudal centrum,
incomplete ~twenty-seventh caudal centrum, ilial fragment, two pubic fragments,
distal ischium, proximal tibia, partial metatarsal I, metatarsal II (335 mm),
phalanx II-1 (130 mm), phalanx II-2 (80 mm), incomplete metatarsal III, phalanx
III-1 fragment, phalanx III-2 (80 mm), phalanx III-3, incomplete metatarsal
IV, phalanx IV-1 (80 mm), phalanx IV-2, phalanx IV-3 (60 mm), partial pedal
ungual IV (Dalman et al., 2012; described in Dalman, 2014a)
?(YPM 57726; paratype of Allosaurus lucasi) ?dentary fragment, angular
fragment (Dalman, 2014a)
ribs, eighteen caudal vertebrae, scapula, metatarsal (Holt, 1940)
fragmentary skeletons including teeth (Armstrong et al., 1987)
tooth (Bollan, 1991)
tooth (Fiorillo and May, 1996)
partial skeleton (Schumacher and Liggett, 2004)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Montana, US (Rattlesnake
Ridge Quarry)
(ANS 21123; "Urinator montanus") (~5.3 m) (mandible ~723 mm) partial
dentaries, partial articular, fifteen teeth (20-31 mm), partial fourteenth dorsal
centrum (80 mm), twenty-three dorsal rib fragments, incomplete first sacral
vertebra (84 mm), fused second and third sacral centra, partial fifth sacral
vertebra, partial proximal caudal centrum, partial scapulae (~460 mm), partial
coracoids, humeri (one incomplete; 293 mm), radii (182, 185 mm), ulnae (218
mm), radiale, intermedium, distal carpal I, distal carpal II, metacarpals I, phalanges
I-1, manual unguals I, metacarpals II, phalanges II-1, phalanges II-2, manual
unguals II, metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3,
manual unguals III, ilial fragment, partial fused pubes (~550 mm), incomplete
ischia (~500 mm), femur (742 mm), proximal tibia (~621 mm), fibulae (one incomplete,
one proximal), partial astragalus, calcaneum, distal metatarsal III (Pirolli,
2004)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, New Mexico, US (Boney Canyon
Quarry, San Ysidro Camarasaur Quarry)
(NMMNH 26071) tooth (Rigby 1982; described by Lucas et al., 1996)
(NMMNH 26073) tooth (Rigby 1982; described by Lucas et al., 1996)
(NMMNH 26074) tooth (Rigby 1982; described by Lucas et al., 1996)
(NMMNH 26083) fourth sacral vertebra, fifth sacral vertebra, first caudal vertebra
(~203 mm), second caudal vertebra (~178 mm), third caudal vertebra (140 mm),
fourth caudal vertebra (140 mm), four chevrons, partial ilium, incomplete ischia,
femora (one incomplete; ~1.1 m), tibia (903 mm), partial fibula, phalanx II-1
(160 mm), phalanx II-2 (122 mm), pedal ungual II, phalanx III-2 (~135 mm) (Williamson
and Chure, 1996)
two vertebrae (Lucas and Hunt, 1985)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Oklahoma, US
(OMNH coll.) material (Langston, 1989)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, South Dakota, US (Fuller's
351)
(SDSM coll.) femur (Bjork, 1983)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Utah, US (Carnegie Quarry,
Cleveland-Lloyd quarry, Green River quarry, Hinkle Site, Rainbow Park)
(11584/BYU VP13019/HEC 464, 488) egg (Hirsch, 1994)
(BMS E25840) (composite) skeleton including humerus (Smith et al., 1999)
(BYU 571/8901; BYU 671/8901 of Eddy and Clarke, 2011; Hinkle specimen) (adult)
incomplete skull (lacking dorsal snout), mandible, atlas, axis, partial scapula,
coracoid, humerus (420 mm), pubis, femur, tibia, metatarsal II (Smith et al.,
1999)
(BYU 2028; "Easter Allosaurus") premaxilla, partial maxilla, nasal, dentary (Lisak, 1980)
(CEU 1719; Al) (8.5 m) partial skeleton including dorsal vertebrae, caudal vertebrae,
coracoid, humeri, ulna, femur, incomplete metatarsal II, pedal ungual II, metatarsal
III, phalanx III-1, phalanx III-2, metatarsal IV (Smith et al., 1999)
(CM 3382) tooth (McIntosh, 1981)
?(CM 3383) vertebrae (McIntosh, 1981)
(CM 3387) teeth, fragments (McIntosh, 1981)
(CM 10002) proximal tibia, proximal fibula (McIntosh, 1981)
?(CM 11843) (juvenile) skull, several vertebral centra, ribs, coracoid, other
elements (McIntosh, 1981)
(CM 11844; = CM 11868; material of Allosaurus "carnegeii")
partial skull, mandible, incomplete skeleton lacking forelimbs and including
femur (842.5 mm), tibia (724 mm) and metatarsal III (360 mm) (McIntosh, 1981)
(CM 21703) skull, presacral vertebrae, caudal vertebrae, ilium, ischium (McIntosh,
1981)
(CM 21705) caudal centrum (McIntosh, 1981)
(CM 21713) ischium, metatarsal, other material (McIntosh, 1981)
(CM 21726) femur (McIntosh, 1981)
(CM 21757) three caudal vertebrae (McIntosh, 1981)
(CM 21769) distal femur (McIntosh, 1981)
(CM 33901) several vertebrae (McIntosh, 1981)
?(CM 33903) gastralia (McIntosh, 1981)
(CM 33957) two caudal vertebrae (McIntosh, 1981)
(CM 33965) two proximal caudal centra, three neural arches, four neural spines
(McIntosh, 1981)
(CM 37004) distal metatarsal (McIntosh, 1981)
(CM 38341) caudal vertebra, ungual (McIntosh, 1981)
(CM 38349) several incomplete metatarsals (McIntosh, 1981)
(CMN 38454) dentary, humerus (Smith et al., 1999)
(DINO 2560, = UUVP 6000; probable intended holotype of "Madsenius trux")
(7.9 m, 1.32 tons) complete skull (845 mm), nearly complete skeleton (lacking
first caudal vertebra, chevrons, forearms, several pedal phalanges) including
femora (880, 850 mm), tibiae (730, 745 mm), astragalus and metatarsals III (375,
372 mm) (Madsen, 1976)
(DINO 3984) femur (605 mm), tibia (55 mm), metatarsal IV (254 mm) (Foster and
Chure, 2006)
(FMNH P1505) (Carrano et al., 2012)
(FMNH P25114) femur (882 mm) (Carrano, 1998)
(LACM 46030) skeleton including partial skull, dentary, distal tarsal IV, metatarsal
II, metatarsal III and metatarsal IV (Smith et al., 2005)
(MCZ 3897) premaxilla, maxilla, cervical vertebrae, dorsal ribs, ilium (398
mm), femur (437 mm), tibia (371 mm) (Madsen, 1976)
(MUP 40CA11) ~10-13th caudal vertebra (Cau and Serventi, 2017)
....(MUP 41CA12 2744) ~9-12th caudal vertebra (Cau and Serventi, 2017)
....(MUP 44CA15 3926) ~14-17th caudal vertebra (Cau and Serventi, 2017)
....(MUP 45CA16) ~14-18th caudal vertebra (Cau and Serventi, 2017)
....(MUP 46CA17) ~16-20th caudal vertebra (Cau and Serventi, 2017)
....(MUP 49CA20 PC69) ~18-21st caudal vertebra (Cau and Serventi, 2017)
....(MUP 52CA23) ~21-24th caudal vertebra (Cau and Serventi, 2017)
....(MUP 61CA32 1484) distal caudal vertebra (Cau and Serventi, 2017)
....(MUP 64CA35 2193) distal caudal vertebra (Cau and Serventi, 2017)
....(MUP 64CA39 PC78) distal caudal vertebra (Cau and Serventi, 2017)
....(MUP 66CA37 407) distal caudal vertebra (Cau and Serventi, 2017)
(ROM 5091) (composite) specimen including maxilla, dentary, humerus, metacarpal
I, metacarpal II, metacarpal III, manual ungual, metatarsals II, metatarsals
III, metatarsals IV (Nicholls and Russell, 1985)
(ROM 12868)
(Carrano et al., 2012)
(UCMP 84804) ungual (UCMP online)
(UDSH C-LQ 004) distal caudal vertebra (Reid, 1990)
(UDSH C-LQ 066) rib (Reid, 1990)
(UDSH C-LQ 068) tibia (Reid, 1990)
(UDSH C-LQ 077) manual unguals (Reid, 1990)
(UDSH C-LQ 109) radius (Reid, 1990)
(UDSH C-LQ 113) pubis (Reid, 1990)
(UDSH C-LQ coll.) 61 elements (Reid, 1990)
(UMNH 1251) premaxilla (Smith et al., 2005)
(UMNH 1765) lacrimal (Carpenter, 2010)
(UMNH 1852) premaxilla (Carpenter, 2010)
(UMNH 3113) (Carrano et al., 2012)
(UMNH 5316) maxilla (Carpenter, 2010)
(UMNH 5326-5328) (Carrano et al., 2012)
(UMNH 5393) maxilla (Carpenter, 2010)
(UMNH 5470) skull (Gates, 2005)
(UMNH 5480) (Carrano et al., 2012)
(UMNH 5754) tooth (?x12.8x? mm) (Heckert et al., 2003)
(UMNH 5812) tooth (22.5x11.5x? mm) (Heckert et al., 2003)
(UMNH 5814) tooth (29.2x13.7x? mm) (Heckert et al., 2003)
(UMNH 5837) tooth (32.9x15.3x? mm) (Heckert et al., 2003)
(UMNH 5918) proximal pubis (Kolb, Davis and Gillette, 1996)
(UMNH 5919) distal scapula (Kolb, Davis and Gillette, 1996)
(UMNH 5920) partial ilium (Kolb, Davis and Gillette, 1996)
(UMNH 5921) fifth sacral rib (Kolb, Davis and Gillette, 1996)
(UMNH 5922) partial eighth or ninth cervical rib (Kolb, Davis and Gillette,
1996)
(UMNH 5923) incomplete fifth sacral vertebra (Kolb, Davis and Gillette, 1996)
(UMNH 5924) sacral vertebra (Kolb, Davis and Gillette, 1996)
(UMNH 5925) partial distal chevron (Kolb, Davis and Gillette, 1996)
(UMNH 5926) mid chevron (145 mm) (Kolb, Davis and Gillette, 1996)
(UMNH 6052) premaxilla (Carpenter, 2010)
(UMNH 6100) tooth (24.8x11.9x? mm) (Heckert et al., 2003)
(UMNH 6140) tooth (36x15x? mm) (Heckert et al., 2003)
(UMNH 6142) tooth (42.8x20.6x? mm) (Heckert et al., 2003)
(UMNH 6145) tooth (23x11.1x? mm) (Heckert et al., 2003)
(UMNH 6153) tooth (41.5x18.5x? mm) (Heckert et al., 2003)
(UMNH 6233) tooth (38.6x17x? mm) (Heckert et al., 2003)
(UMNH 6234) tooth (21.5x12.3x? mm) (Heckert et al., 2003)
(UMNH 6237) tooth (35.3x17.1x? mm) (Heckert et al., 2003)
(UMNH 6242) tooth (41.5x19.6x? mm) (Heckert et al., 2003)
(UMNH 6317) (Carrano et al., 2012)
(UMNH 6340) (Carrano et al., 2012)
(UMNH 6363) maxilla (Carpenter, 2010)
(UMNH 6365) (Carrano et al., 2012)
(UMNH 6400) (Carrano et al., 2012)
(UMNH 6408) (Carrano et al., 2012)
(UMNH 6473) (Carrano et al., 2012)
(UMNH 6475) (Carrano et al., 2012)
(UMNH 6499) (Carrano et al., 2012)
(UMNH 6500) premaxilla (Carpenter, 2010)
(UMNH 6502) (Carrano et al., 2012)
(UMNH 6504) premaxilla (Carpenter, 2010)
(UMNH 6505) premaxilla (Carpenter, 2010)
(UMNH 7190) (Carrano et al., 2012)
(UMNH 7408) (Carrano et al., 2012)
(UMNH 7411) (Carrano et al., 2012)
(UMNH 7442) tooth (31.6x15.7x? mm) (Heckert et al., 2003)
(UMNH 7445) tooth (35.1x18.9x? mm) (Heckert et al., 2003)
(UMNH 7785) lacrimal (Carpenter, 2010)
(UMNH 7786) lacrimal (Carpenter, 2010)
(UMNH 7794) (Carrano et al., 2012)
(UMNH 7880) (Carrano et al., 2012)
(UMNH 7882) (Carrano et al., 2012)
(UMNH 7884) (Carrano et al., 2012)
(UMNH 7885) (Carrano et al., 2012)
(UMNH 7889-7891) (Carrano et al., 2012)
(UMNH 7895) (Carrano et al., 2012)
(UMNH 7898) (Carrano et al., 2012)
(UMNH 7908) (Carrano et al., 2012)
(UMNH 7922) (Carrano et al., 2012)
(UMNH 7926-7930) (Carrano et al., 2012)
(UMNH 7932) (Carrano et al., 2012)
(UMNH 7934) (Carrano et al., 2012)
(UMNH 7937) (Carrano et al., 2012)
(UMNH 7938) (Carrano et al., 2012)
(UMNH 7957) (Carrano et al., 2012)
(UMNH 7966) (Carrano et al., 2012)
(UMNH 8098) lacrimal (Carpenter, 2010)
(UMNH 8102) (Carrano et al., 2012)
(UMNH 8123) (Carrano et al., 2012)
(UMNH 8142) (Carrano et al., 2012)
(UMNH 8151) (Carrano et al., 2012)
(UMNH 8229) (Carrano et al., 2012)
(UMNH 8240) (Carrano et al., 2012)
(UMNH 8241) (Carrano et al., 2012)
(UMNH 8258) premaxilla (Carpenter, 2010)
(UMNH 8355) (Carrano et al., 2012)
(UMNH 8358) posterior cervical vertebra (Evers, Rauhut, Milner, McFeeters and
Allain, 2015)
(UMNH 8365) posterior cervical vertebra (Evers, Rauhut, Milner, McFeeters and
Allain, 2015)
(UMNH 8397) (Carrano et al., 2012)
(UMNH 8475) jugal (Carpenter, 2010)
(UMNH 8484) (Carrano et al., 2012)
(UMNH 8488) posterior cervical vertebra (Evers, Rauhut, Milner, McFeeters and
Allain, 2015)
(UMNH 8489) posterior cervical vertebra (Evers, Rauhut, Milner, McFeeters and
Allain, 2015)
(UMNH 8971) lacrimal (Carpenter, 2010)
(UMNH 8972) jugal (Carpenter, 2010)
(UMNH 8974) jugal (Carpenter, 2010)
(UMNH 8975) jugal (Carpenter, 2010)
(UMNH 8976) jugal (Carpenter, 2010)
(UMNH 9083) jugal (Carpenter, 2010)
(UMNH 9084) jugal (Carpenter, 2010)
(UMNH 9085) jugal (Carpenter, 2010)
(UMNH 9086) jugal (Carpenter, 2010)
(UMNH 9087) jugal (Carpenter, 2010)
(UMNH 9088) jugal (Chure and Loewen, 2020)
(UMNH 9090) incomplete postorbital (Carpenter, 2010)
(UMNH 9096) postorbital (Carpenter, 2010)
(UMNH 9097) postorbital (Carpenter, 2010)
(UMNH 9099) postorbital (Carpenter, 2010)
(UMNH 9100) postorbital (Carpenter, 2010)
(UMNH 9103) postorbital (Carpenter, 2010)
(UMNH 9106) postorbital (Carpenter, 2010)
(UMNH 9108) postorbital (Carpenter, 2010)
(UMNH 9112) postorbital (Carpenter, 2010)
(UMNH 9147) (Carrano et al., 2012)
(UMNH 9149) (Carrano et al., 2012)
(UMNH 9160) premaxilla (Carpenter, 2010)
(UMNH 9162) (Carrano et al., 2012)
(UMNH 9168) (Carrano et al., 2012)
(UMNH 9169) (different sizes; one is a typo?) two maxillae (Carpenter, 2010)
(UMNH 9180) (Carrano et al., 2012)
(UMNH 9191) (Carrano et al., 2012)
(UMNH 9201) maxilla (Carpenter, 2010)
(UMNH 9210) incomplete maxilla (Carpenter, 2010)
(UMNH 9211) maxilla (Smith et al., 2005)
(UMNH 9212) (Carrano et al., 2012)
(UMNH 9215) maxilla (Carpenter, 2010)
(UMNH 9218) maxilla (Smith et al., 2005)
(UMNH 9224) incomplete lacrimal (Carpenter, 2010)
(UMNH 9229) maxilla (Carpenter, 2010)
(UMNH 9231) premaxilla (Carpenter, 2010)
(UMNH 9241) premaxilla (Carpenter, 2010)
(UMNH 9264) premaxilla (Carpenter, 2010)
(UMNH 9348) premaxilla (Carpenter, 2010)
(UMNH 9252) premaxilla (Carpenter, 2010)
(UMNH 9261) incomplete premaxilla (Carpenter, 2010)
(UMNH 9262) premaxilla (Carpenter, 2010)
(UMNH 9263) premaxilla (Carpenter, 2010)
(UMNH 9265) premaxilla (Carpenter, 2010)
(UMNH 9270) incomplete premaxilla (Carpenter, 2010)
(UMNH 9272) maxilla (Carpenter, 2010)
(UMNH 9273) maxilla (Smith et al., 2005)
(UMNH 9274) maxilla (Carpenter, 2010)
(UMNH 9275) maxilla (Smith et al., 2005)
(UMNH 9323) (Carrano et al., 2012)
(UMNH 9327) (Carrano et al., 2012)
(UMNH 9365) dentary (Smith et al., 2005)
(UMNH 9366) (Carrano et al., 2012)
(UMNH 9369) dentary (Smith et al., 2005)
(UMNH 9376) (Carrano et al., 2012)
(UMNH 9401) (Carrano et al., 2012)
(UMNH 9469) incomplete lacrimal (Carpenter, 2010)
(UMNH 9470) (Carrano et al., 2012)
(UMNH 9471) lacrimal (Carpenter, 2010)
(UMNH 9472) lacrimal (Carpenter, 2010)
(UMNH 9473) incomplete lacrimal (Carpenter, 2010)
(UMNH 9474) lacrimal (Carpenter, 2010)
(UMNH 9475) lacrimal (Carpenter, 2010)
(UMNH 9476) incomplete lacrimal (Carpenter, 2010)
(UMNH 9478) lacrimal (Carpenter, 2010)
(UMNH 9479) incomplete lacrimal (Carpenter, 2010)
(UMNH 9480) (Carrano et al., 2012)
(UMNH 9500) (Carrano et al., 2012)
(UMNH 9502) (Carrano et al., 2012)
(UMNH 9505) (Carrano et al., 2012)
(UMNH 9514) (Carrano et al., 2012)
(UMNH 9528) jugal (Chure and Loewen, 2020)
(UMNH 9570) incomplete jugal (Chure and Loewen, 2020)
(UMNH 9709) (Carrano et al., 2012)
(UMNH 10192) posterior cervical vertebra (Evers, Rauhut, Milner, McFeeters and
Allain, 2015)
(UMNH 10360) (Carrano et al., 2012)
(UMNH 10386) (Carrano et al., 2012)
(UMNH 10393) premaxilla (Carpenter, 2010)
(UMNH 10779) (Carrano et al., 2012)
(UMNH 10781; = UUVP 3811) first caudal vertebra (115 mm) (Carpenter, Sanders,
McWhinney and Wood, 2005)
(UMNH 11031) (Carrano et al., 2012)
(UMNH 11463) (Carrano et al., 2012)
(UMNH 11531) lacrimal (Carpenter, 2010)
(UMNH 12000) jugal (Carpenter, 2010)
(UMNH 12005) lacrimal (Carpenter, 2010)
(UMNH 12042) partial pelvis (Gates, 2005)
(UMNH 12176) lacrimal (Carpenter, 2010)
(UMNH 12193) partial pelvis (Gates, 2005)
(UMNH 12194) partial pelvis (Gates, 2005)
(UMNH 12196) femur (Gates, 2005)
(UMNH 12197) tibia (Gates, 2005)
(UMNH 12226) skull (Gates, 2005)
(UMNH 12231) (Carrano et al., 2012)
(UMNH 16584) (Carrano et al., 2012)
(UMNH 16585) (Carrano et al., 2012)
(UMNH 16659) maxilla (Carpenter, 2010)
(UMNH 16664) lacrimal (Carpenter, 2010)
(UMNH 16665) lacrimal (Carpenter, 2010)
(UMNH 89088) jugal (Carpenter, 2010)
(UMNH CLQ-03-34) maxilla (Carpenter, 2010)
(UUVP 3) (Molnar, 1991)
(UUVP 10-245) premaxilla (Madsen, 1976)
(UUVP 30) humerus (Smith et al., 1999)
(UUVP 30-16) femur (535 mm) (Madsen, 1976)
(UUVP 30-17) femur (605 mm) (Madsen, 1976)
(UUVP 30-35) femur (465 mm) (Madsen, 1976)
(UUVP 30-55) tibia (343 mm) (Madsen, 1976)
(UUVP 30-76) ulna (Hanna, 2002)
(UUVP 30-77) humerus (202 mm) (Madsen, 1976)
(UUVP 30-90) metatarsal III (300 mm) (Madsen, 1976)
(UUVP 30-293) premaxilla (Madsen, 1976)
(UUVP 30-296) dentary (Madsen, 1976)
(UUVP 30-714) tibia (500 mm) (Madsen, 1976)
(UUVP 30-723) premaxilla (Madsen, 1976)
(UUVP 30-724) femur (525 mm) (Madsen, 1976)
(UUVP 30-743) femur (535 mm) (Madsen, 1976)
(UUVP 30-752) femur (465 mm) (Madsen, 1976)
(UUVP 30-778) humerus (280 mm) (Peterson, Isakson and Madsen, 1972)
(UUVP 30-782) metatarsal III (318 mm) (Madsen, 1976)
(UUVP 30-783) metatarsal IV (Hanna, 2002)
(UUVP 33) basicranium (Chure and Madsen, 1996)
(UUVP 40) basicranium, pedal phalanx III-1 (Madsen, 1976; Chure and Madsen,
1996)
(UUVP 40-21) astragalus (Madsen, 1976)
(UUVP 40-22) astragalus (Madsen, 1976)
(UUVP 40-25) astragalus (Madsen, 1976)
(UUVP 40-214) metatarsal III (333 mm) (Madsen, 1976)
(UUVP 40-216) metatarsal III (296 mm) (Madsen, 1976)
(UUVP 40-217) metatarsal III (285 mm) (Madsen, 1976)
(UUVP 40-219) metatarsal III (269 mm) (Madsen, 1976)
(UUVP 40-221) metatarsal III (247 mm) (Madsen, 1976)
(UUVP 40-222) metatarsal III (243 mm) (Madsen, 1976)
(UUVP 40-227) humerus (252 mm) (Madsen, 1976)
(UUVP 40-268) femur (535 mm) (Madsen, 1976)
(UUVP 40-297) tibia (612 mm) (Madsen, 1976)
(UUVP 40-298) tibia (525 mm) (Madsen, 1976)
(UUVP 40-301) tibia (552 mm) (Madsen, 1976)
(UUVP 40-304) tibia (552 mm) (Madsen, 1976)
(UUVP 40-306) tibia (525 mm) (Madsen, 1976)
(UUVP 40-308) tibia (540 mm), astragalus (Madsen, 1976)
(UUVP 40-331) dentary (Madsen, 1976)
(UUVP 40-453) fifth sacral vertebra (Kolb, Davis and Gillette, 1996)
(UUVP 40-553) dentary (Madsen, 1976)
(UUVP 40-586) maxilla (Madsen, 1976)
(UUVP 40-587) maxilla (Madsen, 1976)
(UUVP 40-601) premaxilla (Madsen, 1976)
(UUVP 40-603) premaxilla (Madsen, 1976)
(UUVP 40-604) premaxilla (Madsen, 1976)
(UUVP 40-722) premaxilla (Madsen, 1976)
(UUVP 40-729) humerus (240 mm) (Madsen, 1976)
(UUVP 40-768) tibia (475 mm) (Madsen, 1976)
(UUVP 71-1) (Molnar, 1991)
(UUVP 71-3) (Molnar, 1991)
(UUVP 71-151) (Molnar, 1991)
(UUVP 86) radius (Madsen, 1976)
(UUVP 139) premaxilla (Madsen, 1976)
(UUVP 142) dentary (Madsen, 1976)
(UUVP 154) tibia (410 mm) (Madsen, 1976)
(UUVP 169) ilium (Kolb, Davis and Gillette, 1996)
(UUVP 177) two distal caudal vertebrae (Peterson, Isakson and Madsen, 1972)
(UUVP 181) pubis (Kolb, Davis and Gillette, 1996)
(UUVP 200) dentary (Smith et al., 1999)
(UUVP 249) tibia (542 mm) (Madsen, 1976)
(UUVP 273) humerus (290 mm) (Madsen, 1976)
(UUVP 294) basicranium (Chure and Madsen, 1996)
(UUVP 387) partial furcula (Chure and Madsen, 1996)
(UUVP 454) astragalus (Madsen, 1976)
(UUVP 492) femur (480 mm) (Madsen, 1976)
(UUVP 566) humerus (199 mm) (Madsen, 1976)
(UUVP 652) postorbital (Smith et al., 1999)
(UUVP 669) mid chevron (143 mm) (Kolb, Davis and Gillette, 1996)
(UUVP 687) radius (Madsen, 1976)
(UUVP 699) dentary (Madsen, 1976)
(UUVP 702) dentary (Madsen, 1976)
(UUVP 718) femur (535 mm) (Madsen, 1976)
(UUVP 740) premaxilla (Madsen, 1976)
(UUVP 778) humerus (Smith et al., 1999)
(UUVP 837) chevron (Peterson, Isakson and Madsen, 1972)
(UUVP 847) femur (450 mm) (Madsen, 1976)
(UUVP 856) premaxilla (Madsen, 1976)
(UUVP 870) dentary (Madsen, 1976)
(UUVP 871) dentary (Madsen, 1976)
(UUVP 915) distal chevron (69 mm) (Kolb, Davis and Gillette, 1996)
(UUVP 919) pubis (Kolb, Davis and Gillette, 1996)
(UUVP 1010) proximal caudal vertebra (Hanna, 2002)
(UUVP 1046) dentary (Madsen, 1976)
(UUVP 1075) metatarsal III (284 mm) (Madsen, 1976)
(UUVP 1086) premaxilla (Madsen, 1976)
(UUVP 1165) femur (865 mm) (Madsen, 1976)
(UUVP 1169) humerus (379 mm) (Madsen, 1976)
(UUVP 1233) maxilla (Madsen, 1976)
(UUVP 1334) humerus (334 mm) (Madsen, 1976)
(UUVP 1364) femur (575 mm) (Madsen, 1976)
(UUVP 1365) maxilla (Madsen, 1976)
(UUVP 1403) jugal (Smith et al., 1999)
(UUVP 1414) epipterygoid (Eddy and Clarke, 2011)
(UUVP 1473) tibia (637 mm) (Madsen, 1976)
(UUVP 1517) maxilla (Madsen, 1976)
(UUVP 1528) scapula (Peterson, Isakson and Madsen, 1972)
(UUVP 1551) metatarsal III (367 mm) (Madsen, 1976)
(UUVP 1582) maxilla (Madsen, 1976)
(UUVP 1622) premaxilla (Madsen, 1976)
(UUVP 1657) pedal phalanx III-1 (Madsen, 1976)
(UUVP 1685) postorbital (Smith et al., 1999)
(UUVP 1743) metatarsal III (313 mm) (Madsen, 1976)
(UUVP 1847) posterior dorsal rib (Peterson, Isakson and Madsen, 1972)
(UUVP 1848) phalanx (Peterson, Isakson and Madsen, 1972)
(UUVP 1849) three distal caudal vertebrae (Peterson, Isakson and Madsen, 1972)
(UUVP 1850) caudal vertebrae (Peterson, Isakson and Madsen, 1972)
(UUVP 1851) pedal phalanx IV-1 (Peterson, Isakson and Madsen, 1972)
(UUVP 1852) premaxilla (Peterson, Isakson and Madsen, 1972)
(UUVP 1853) pedal ungual II (Peterson, Isakson and Madsen, 1972)
(UUVP 1854) metacarpal (Peterson, Isakson and Madsen, 1972)
(UUVP 1855) metacarpal (Peterson, Isakson and Madsen, 1972)
(UUVP 1856-1858) bone (Peterson, Isakson and Madsen, 1972)
(UUVP 1862) postorbital (Smith et al., 1999)
(UUVP 1863) premaxilla (Madsen, 1976)
(UUVP 1865) premaxilla (Madsen, 1976)
(UUVP 1866) premaxilla (Madsen, 1976)
(UUVP 1869) premaxilla (Madsen, 1976)
(UUVP 1872) premaxilla (Madsen, 1976)
(UUVP 1873) premaxilla (Madsen, 1976)
(UUVP 1875) premaxilla (Madsen, 1976)
(UUVP 1876) premaxilla (Madsen, 1976)
(UUVP 1878) premaxilla (Madsen, 1976)
(UUVP 1879) premaxilla (Madsen, 1976)
(UUVP 1880) maxilla (Madsen, 1976)
(UUVP 1881) maxilla (Madsen, 1976)
(UUVP 1883) maxilla (Madsen, 1976)
(UUVP 1884) maxilla (Madsen, 1976)
(UUVP 1886) maxilla (Madsen, 1976)
(UUVP 1887) maxilla (Madsen, 1976)
(UUVP 1889) maxilla (Madsen, 1976)
(UUVP 1890) maxilla (Madsen, 1976)
(UUVP 1892) maxilla (Madsen, 1976)
(UUVP 1893) maxilla (Madsen, 1976)
(UUVP 1894) maxilla (Madsen, 1976)
(UUVP 1895) dentary (Madsen, 1976)
(UUVP 1896) dentary (Madsen, 1976)
(UUVP 1897) dentary (Madsen, 1976)
(UUVP 1898) dentary (Madsen, 1976)
(UUVP 1900) dentary (Madsen, 1976)
(UUVP 1901) dentary (Madsen, 1976)
(UUVP 1903) dentary (Madsen, 1976)
(UUVP 1904) dentary (Madsen, 1976)
(UUVP 1905) dentary (Madsen, 1976)
(UUVP 1906) dentary (Madsen, 1976)
(UUVP 1907) dentary (Madsen, 1976)
(UUVP 1908) dentary (Madsen, 1976)
(UUVP 1909) dentary (Madsen, 1976)
(UUVP 1910) dentary (Madsen, 1976)
(UUVP 1927) premaxilla (Madsen, 1976)
(UUVP 1934) postorbital (Smith et al., 1999)
(UUVP 1936) postorbital (Smith et al., 1999)
(UUVP 1945) premaxilla (Madsen, 1976)
(UUVP 1980) humerus (156 mm) (Madsen, 1976)
(UUVP 1981) humerus (162 mm) (Madsen, 1976)
(UUVP 1989) metatarsal III (228 mm) (Madsen, 1976)
(UUVP 1991) premaxilla (Madsen, 1976)
(UUVP 2001) dentary (Madsen)
(UUVP 2067) basicranium (Chure and Madsen, 1996)
(UUVP 2175) postorbital (Smith et al., 1999)
(UUVP 2186) maxilla (Madsen, 1976)
(UUVP 2226) fifth sacral vertebra (Kolb, Davis and Gillette, 1996)
(UUVP 2252) cervical rib (Peterson, Isakson and Madsen, 1972)
(UUVP 2280) femur (505 mm) (Madsen, 1976)
(UUVP 2327) humerus (244 mm) (Madsen, 1976)
(UUVP 2350) distal chevron (64 mm) (Kolb, Davis and Gillette, 1996)
(UUVP 2456) dentary (Madsen, 1976)
(UUVP 2545) premaxilla (Madsen, 1976)
(UUVP 2549) astragalus (Madsen, 1976)
(UUVP 2550) mid chevron (107 mm) (Kolb, Davis and Gillette, 1996)
(UUVP 2559) femur (435 mm) (Madsen, 1976)
(UUVP 2567) astragalus (Madsen, 1976)
(UUVP 2600) premaxilla (Madsen, 1976)
(UUVP 2753) dorsal rib (Hanna, 2002)
(UUVP 2758) postorbital (Smith et al., 1999)
(UUVP 2765) humerus (251 mm) (Madsen, 1976)
(UUVP 2843) premaxilla (Madsen, 1976)
(UUVP 2850) basicranium (Chure and Madsen, 1996)
(UUVP 2887) maxilla (Madsen, 1976)
(UUVP 2903) dentary (Madsen, 1976)
(UUVP 2923) metatarsal III (284 mm) (Madsen, 1976)
(UUVP 2939) pedal phalanx (Hanna, 2002)
(UUVP 2997) phalanx (Hanna, 2002)
(UUVP 3036) premaxilla (Madsen, 1976)
(UUVP 3082) quadrate, braincase (Eddy and Clarke, 2011)
(UUVP 3133) seventh cervical rib (Kolb, Davis and Gillette, 1996)
(UUVP 3164) femur (475 mm) (Madsen, 1976)
(UUVP 3192) femur (545 mm) (Madsen, 1976)
(UUVP 3203) basicranium (Chure and Madsen, 1996)
(UUVP 3243) ischium (Peterson, Isakson and Madsen, 1972)
(UUVP 3249) tibia (543 mm) (Madsen, 1976)
(UUVP 3279) maxilla (Madsen, 1976)
(UUVP 3287) basicranium (Chure and Madsen, 1996)
(UUVP 3304) basicranium (Chure and Madsen, 1996)
(UUVP 3312) humerus (183 mm) (Madsen, 1976)
(UUVP 3342) humerus (255 mm) (Madsen, 1976)
(UUVP 3385) femur ( mm) (Madsen, 1976)
(UUVP 3389) dentary (Madsen, 1976)
(UUVP 3435) humerus (343 mm) (Peterson, Isakson and Madsen, 1972)
(UUVP 3529) premaxilla (Madsen, 1976)
(UUVP 3587) maxilla (Madsen, 1976)
(UUVP 3607) humerus (386 mm) (Madsen, 1976)
(UUVP 3630) metatarsal III (191 mm) (Madsen, 1976)
(UUVP 3638) maxilla (Madsen, 1976)
(UUVP 3670) premaxilla (Madsen, 1976)
(UUVP 3694) femur (905 mm) (Madsen, 1976)
(UUVP 3724) premaxilla (Madsen, 1976)
(UUVP 3733) astragalus (Madsen, 1976)
(UUVP 3753) astragalus (Madsen, 1976)
(UUVP 3758) postorbital (Smith et al., 1999)
(UUVP 3771) two mid caudal vertebrae, chevron (Madsen, 1976)
(UUVP 3773) chevron (Peterson, Isakson and Madsen, 1972)
(UUVP 3810) dentary (Madsen, 1976)
(UUVP 3811) dorsal vertebra (Hanna, 2002)
(UUVP 3833) tibia (553 mm) (Madsen, 1976)
(UUVP 3835) tibia (540 mm) (Madsen, 1976)
(UUVP 3836) maxilla (Madsen, 1976)
(UUVP 3943) fifth sacral rib (Kolb, Davis and Gillette, 1996)
(UUVP 3980) femur (630 mm) (Madsen, 1976)
(UUVP 3995) premaxilla (Madsen, 1976)
(UUVP 4029) dentary (Madsen, 1976)
(UUVP 4122) postorbital (Smith et al., 1999)
(UUVP 4127) astragalus (Madsen, 1976)
(UUVP 4159) phalanx (Peterson, Isakson and Madsen, 1972)
(UUVP 4201) bone (Peterson, Isakson and Madsen, 1972)
(UUVP 4213) maxilla (Madsen, 1976)
(UUVP 4220) humerus (150 mm) (Madsen, 1976)
(UUVP 4223) metatarsal III (345 mm) (Madsen, 1976)
(UUVP 4233) dentary (Madsen, 1976)
(UUVP 4320) chevron (Peterson, Isakson and Madsen, 1972)
(UUVP 4385) metatarsal III (338 mm) (Madsen, 1976)
(UUVP 4387) humerus (254 mm) (Madsen, 1976)
(UUVP 4445) metatarsal III (292 mm) (Madsen, 1976)
(UUVP 4493) astragalus (Madsen, 1976)
(UUVP 4556) postorbital (Smith et al., 1999)
(UUVP 4596) premaxilla (Madsen, 1976)
(UUVP 4674) postorbital (Smith et al., 1999)
(UUVP 4778) dentary (Madsen, 1976)
(UUVP 4779) maxilla (Madsen, 1976)
(UUVP 4780) maxilla (Madsen, 1976)
(UUVP 4785) metatarsal III (336 mm) (Madsen, 1976)
(UUVP 4792) humerus (264 mm) (Madsen, 1976)
(UUVP 4813) maxilla (Madsen, 1976)
(UUVP 4877) metatarsal III (280 mm) (Madsen, 1976)
(UUVP 4895) caudal vertebrae (Peterson, Isakson and Madsen, 1972)
(UUVP 4908) humerus (280 mm) (Madsen, 1976)
(UUVP 4946) rib (Hanna, 2002)
(UUVP 4958) metatarsal III (298 mm) (Madsen, 1976)
(UUVP 5079) dentary (Madsen, 1976)
(UUVP 5141) astragalus (Madsen, 1976)
(UUVP 5160) postorbital (Smith et al., 1999)
(UUVP 5186) mid chevron (175 mm) (Kolb, Davis and Gillette, 1996)
(UUVP 5198) lacrimal (Eddy and Clarke, 2011)
(UUVP 5237) metatarsal III (291 mm) (Madsen, 1976)
(UUVP 5256) two caudal vertebrae, chevron (Hanna, 2002)
(UUVP 5289) dentary (Madsen, 1976)
(UUVP 5300) (female) tibia (622 mm) (Madsen, 1976)
(UUVP 5301) tibia (567 mm) (Madsen, 1976)
(UUVP 5302) femur (695 mm) (Madsen, 1976)
(UUVP 5315) premaxilla (Madsen, 1976)
(UUVP 5346) basicranium (Chure and Madsen, 1996)
(UUVP 5358) astragalus (Madsen, 1976)
(UUVP 5361) tibia (596 mm) (Madsen, 1976)
(UUVP 5379) metatarsal III (257 mm) (Madsen, 1976)
(UUVP 5386) maxilla (Madsen, 1976)
(UUVP 5391) pubis (Kolb, Davis and Gillette, 1996)
(UUVP 5410) fifth sacral rib (Kolb, Davis and Gillette, 1996)
(UUVP 5415) metatarsal III (285 mm) (Madsen, 1976)
(UUVP 5420) astragalus (Madsen, 1976)
(UUVP 5423) humerus (205 mm) (Madsen, 1976)
(UUVP 5427) premaxilla, maxilla, dentary, fifth sacral vertebra (Madsen, 1976;
Kolb, Davis and Gillette, 1996)
(UUVP 5436) astragalus (Madsen, 1976)
(UUVP 5445) metatarsal III (310 mm) (Madsen, 1976)
(UUVP 5472) humerus (351 mm) (Madsen, 1976)
(UUVP 5490) premaxilla (Madsen, 1976)
(UUVP 5496) humerus (276 mm) (Madsen, 1976)
(UUVP 5499) maxilla (Madsen, 1976)
(UUVP 5501) humerus (295 mm) (Madsen, 1976)
(UUVP 5566) premaxilla (Madsen, 1976)
(UUVP 5577) humerus (333 mm) (Madsen, 1976)
(UUVP 5579) metatarsal III (252 mm) (Madsen, 1976)
(UUVP 5582) postorbital (Smith et al., 1999)
(UUVP 5583) basicranium (Chure and Madsen, 1996)
(UUVP 5599) scapula (Peterson, Isakson and Madsen, 1972)
(UUVP 5626) chevron (Peterson, Isakson and Madsen, 1972)
(UUVP 5658) distal caudal vertebra (Hanna, 2002)
(UUVP 5659) distal caudal vertebra (Hanna, 2002)
(UUVP 5660) rib (Hanna, 2002)
(UUVP 5661) rib (Hanna, 2002)
(UUVP 5669) pedal phalanx IV-2 (Hanna, 2002)
(UUVP 5748) maxillae, pterygoid, dentary (Madsen, 1976; Smith et al., 1999)
(UUVP 5753) furcula (Chure and Madsen, 1996)
(UUVP 5748) basicranium (Chure and Madsen, 1996)
(UUVP 5754) furcula (Chure and Madsen, 1996)
(UUVP 5843) basicranium (Chure and Madsen, 1996)
(UUVP 5849) basicranium (Chure and Madsen, 1996)
(UUVP 5942) basicranium (Chure and Madsen, 1996)
(UUVP 5943) basicranium (Chure and Madsen, 1996)
(UUVP 5958) postorbital (Eddy and Clarke, 2011)
(UUVP 5961) braincase (Eddy and Clarke, 2011)
(UUVP 5969) basicranium (Chure and Madsen, 1996)
(UUVP 5983) metatarsal III (276 mm) (Madsen, 1976)
(UUVP 5985) ilium, ischium, metatarsal III (350 mm) (Madsen, 1976)
(UUVP 5988) tibia (577 mm) (Madsen, 1976)
(UUVP 5990) tibia (546 mm) (Madsen, 1976)
(UUVP 5991) femur (555 mm) (Madsen, 1976)
(UUVP 5993) femur (800 mm) (Madsen, 1976)
(UUVP 5998) astragalus (Madsen, 1976)
(UUVP 6023) scapula, femur (245 mm) (Madsen, 1976)
(UUVP 6061) astragalus (Madsen, 1976)
(UUVP 6062) astragalus (Madsen, 1976)
(UUVP 6063) astragalus (Madsen, 1976)
(UUVP 6100) partial furcula (Chure and Madsen, 1996)
(UUVP 6101) furcula (Chure and Madsen, 1996)
(UUVP 6102) partial furcula (Chure and Madsen, 1996)
(UUVP 6103) astragalus (Madsen, 1976)
(UUVP 6132) partial furcula (Chure and Madsen, 1996)
(UUVP 6625) proximal caudal vertebra (Madsen, 1976)
(UUVP 6737) premaxilla (Madsen, 1976)
(UUVP 6738) astragalus (Madsen, 1976)
(UUVP 6740) premaxilla (Madsen, 1976)
(UUVP 6742) dentary (Madsen, 1976)
(UUVP 6788) pedal phalanx III-1 (Hanna, 2002)
(UUVP 6912) basicranium (Chure and Madsen, 1996)
(UUVP 6979) pubis (Kolb, Davis and Gillette, 1996)
(UUVP 7145) braincase (Eddy and Clarke, 2011)
(UUVP 7163) ilium (Kolb, Davis and Gillette, 1996)
(UUVP 10016) ilium (Kolb, Davis and Gillette, 1996)
(UUVP 10093) dentary (Madsen, 1976)
(UUVP 10111) postorbital (Smith et al., 1999)
(UUVP 10136) cervical vertebra (Hanna, 2002)
(UUVP 10149) astragalus (Madsen, 1976)
(UUVP 10154) humerus (340 mm) (Madsen, 1976)
(UUVP 10161) humerus (280 mm) (Madsen, 1976)
(UUVP 10169) metatarsal III (365 mm) (Madsen, 1976)
(UUVP 10173) jugal (Smith et al., 1999)
(UUVP 10220) pedal phalanx II-1 (Hanna, 2002)
(UUVP 10237) maxilla (Madsen, 1976)
(UUVP 10248) tibia (695 mm) (Madsen, 1976)
(UUVP 10249) astragalus (Madsen, 1976)
(UUVP 10250) dentary (Madsen, 1976)
(UUVP 10863) ilium (Malafaia et al., 2017)
(UUVP 10908) pedal phalanx IV-1 (Hanna, 2002)
(UUVP 11497) scapula (Kolb, Davis and Gillette, 1996)
(UUVP 11498) scapula (Kolb, Davis and Gillette, 1996)
(UUVP 11499) ilium (Kolb, Davis and Gillette, 1996)
(UUVP 11500) ilium (Kolb, Davis and Gillette, 1996)
(UUVP 11690) furcula (Chure and Madsen, 1996)
(UUVP 16645) braincase (Eddy and Clarke, 2011)
(UUVP 40607) postorbital (Smith et al., 1999)
(UUVP 40609) postorbital (Smith et al., 1999)
(UUVP 40610) postorbital (Smith et al., 1999)
(UUVP 40-738) metatarsal III (343 mm) (Madsen, 1976)
(UUVP 40-739) metatarsal III (298 mm) (Madsen, 1976)
(UUVP A1-1) (Molnar, 1991)
(UUVP Q-6) (Molnar, 1991)
(UUVP Q-19) (Molnar, 1991)
(UUVP X31) (Molnar, 1991)
(UUVP coll.) rib (Hanna, 2002)
(YPM 4944) (composite) skeleton including premaxilla, humerus (335 mm), ulna,
metacarpal I, metacarpal II, metacarpal III, ilium, pubis, femur (526 mm), tibia
(450 mm), astragalus, phalanx II-1, phalanx II-2, pedal ungual II and metatarsal
III (238 mm) (Ostrom, 1969)
(YPM 54376) tooth (YPM online)
(YPM 55898) two fragmentary teeth (Marsh, 1871)
(YPM PU 14554; AMNH 14554 of Eddy and Clarke, 2011) skeleton including maxilla,
lacrimal, nasal, squamosal, mandible, femora (780, 786 mm) (Witmer, 1997)
(YPM PU 14554 A) ilium (YPM online)
(YPM PU 14554 B) pubes (YPM online)
(YPM PU 14554 C) fibula (YPM online)
(YPM PU 14554 D) femora (YPM online)
(YPM PU 14554 E) tibia, fibula, astragalus, calcaneum (YPM online)
(YPM PU 14554 F) tibiae (YPM online)
(YPM PU 14554 G; = PU 11 and PU 12 of Smith et al., 1999?) dentaries (YPM online)
(YPM PU 14554 H; = PU 6 and PU 7 of Smith et al., 1999?) cranial elements (YPM
online)
(YPM PU 14554 I; = PU 3, PU 4 and PU 7 of Smith et al., 1999?) forelimb elements
(YPM online)
(YPM PU 14554 J) hindlimb elements (YPM online)
(YPM PU 14554 K) vertebrae, chevrons (YPM online)
(YPM PU 14554 L) ischium (YPM online)
tooth (Stokes, 1964)
teeth (Chure and Englemann, 1989)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Beside Sauropod Quarry, Nail Quarry,
Reed's Quarry 9, Reed’s Quarry 14, Quarry N, WDC DML quarry)
(AMNH 5759) teeth (AMNH online)
(CM 1254; = CM 1255 in part) premaxilla, two teeth, two sacral vertebrae, humerus,
ischia, four metatarsals, several phalanges (McIntosh, 1981)
(KUVP 1392) centra, scapulae, coracoids, incomplete humerus, radius, manual
ungual I, ilium, femur (Williston, 1901)
(TATE 542) (adult) tooth (36 mm) (Bakker, 1997)
(TATE 543) (adult) tooth (20 mm) (Bakker, 1997)
(TATE 544) (adult) tooth (9.9 mm) (Bakker, 1997)
(TATE 550) (juvenile) tooth (6.6 mm) (Bakker, 1997)
(TATE coll.; material of "Wyomingraptor") 138 juvenile to adult teeth,
three adult individuals including distal caudal vertebrae, forelimb, pubes and
ischia (Bakker, 1997)
(USNM 2323) eight cervical centra, eleven dorsal centra, two sacral centra,
many cervical and dorsal neural processes, ribs, ilium, ischia (490 mm), femur
(645 mm) (Hay, 1908)
(USNM 8302) manual ungual III (Gilmore, 1920)
(WDC BS-749) premaxillary tooth (31.6x10x12 mm) (Diepenbrock, 2020)
(WDC BS-798) premaxillary tooth (11.2x3.9x5.2 mm) (Diepenbrock, 2020)
(WDC BS-846) premaxillary tooth (23.2x5.4x9.4 mm) (Diepenbrock, 2020)
(WDC BS-1777) premaxillary tooth (27x8.9.5x9 mm) (Diepenbrock, 2020)
(WDC DMJ-0026) (Wahl, 2006)
(YPM 1932) proximal caudal vertebra (Marsh, 1884)
Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Colorado, US
(LACM uncatalogued/HEC 629) eggshell (Hirsch, 1994)
(MWC 122.2/HEC 418) eggs, eggshells (Hirsch, 1994)
(MWC 122.2/HEC 462) eggs, eggshells (Hirsch, 1994)
(MWC 122.2/HEC 489) eggs, eggshells (Hirsch, 1994)
(MWC 122.2/HEC 532) eggs, eggshells (Hirsch, 1994)
(MWC 122.3/HEC 457) eggs, eggshells (Hirsch, 1994)
....(MWC 122.3.1/HEC 457; holotype of Prismatoolithus coloradensis) egg
(Hirsch, 1994)
(MWC 122.6) eggs, eggshells (Hirsch, 1994)
(MWC 122.8) eggs, eggshells (Hirsch, 1994)
Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Montana, US (Mother's Day quarry,
O'Hair quarry, Upper Strickland Creek quarry)
(MOR 637) tibia (734 mm), partial pes of eight elements including metatarsal
III (372 mm) (Carrano, 1998)
(MOR 790-8-3-96-142) tooth (MOR online)
specimen (Schimelfening, Woodruff and Norden, 2014)
Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, New Mexico, US (Acoma quarry)
(NMMNH P-38975; private coll.) maxillary fragments, dentary fragments, teeth
(Heckert et al., 2003)
caudal vertebra, partial manual phalanx III-1, ungual (Smith, 1961)
Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Utah, US (DNM 116)
(DINO 11541; holotype of Allosaurus jimmadseni)
(5.6 m, 614 kg, subadult) right half of skull (630 mm), stapes, partial sclerotic
ring, mandible, hyoids, (presacral column 1.814 m) second through eleventh cervical
vertebrae, sixth through eighth cervical ribs, first through twelfth dorsal
vertebrae, second through twelfth dorsal ribs, eighteen rows of gastralia, sacrum
(438 mm), firsth through eighth caudal vertebrae (722 mm), midcaudal vertebra,
sixteen distal caudal vertebrae (991 mm), seventeen chevrons, scapulae, coracoids
(133 mm), furcula, humeri, radius, ulna, carpus, manus, keratinous sheath of
manual ungual I, ilia, pubes, ischia (454 mm), femora (~658 mm), tibiae, fibulae,
astragalus, calcaneum, distal tarsals 3, distal tarsal IV, metatarsal I, metatarsal
II (182 mm), phalanx II-1, phalanx II-2, metatarsal III (225 mm), phalanx III-1,
phalanx III-2, metatarsal IV (195 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal ungual IV (Chure, 2000a)
Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Wyoming, US (Big Al Quarry,
Bone Cabin Quarry, Dana Quarry, Howe Quarry, Lake’s Quarry 1A, Meilyn Quarry,
Poison Creek (Flynn) Quarry-3)
(AMNH 257) femur (920 mm) (Carrano, 1998)
(AMNH 275) femur (910 mm) (Osborn, 1899)
(AMNH 281) femur (AMNH online)
(AMNH 287) fibula (AMNH online)
(AMNH 290) (~9.5 m) partial metacarpal I, femur (985 mm), tibia (810 mm), fibula
(764 mm), astragalus, calcaneum, metatarsal I, phalanx I-1, pedal ungual I,
metatarsal II (375 mm), phalanx II-1, phalanx II-2, pedal ungual II, metatarsal
III (425 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III,
metatarsal IV (360 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4,
pedal ungual IV, metatarsal V (Osborn, 1899)
(AMNH 324) tibia (698 mm), fibula (665 mm), metatarsal I, phalanx I-1, pedal
ungual I, metatarsal II (315 mm), phalanx II-1, phalanx II-2, pedal ungual II,
metatarsal III (352 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal
ungual III, metatarsal IV (330 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal ungual IV, metatarsal V (Osborn, 1899)
(AMNH 393; lost) ten caudal vertebrae, chevron, fragments (AMNH online)
(AMNH 408) ulna, femora (778, 825 mm), tibiae (705, 705 mm), proximal fibula,
metatarsal IV, two phalanges (Carrano, 1998)
(AMNH 468; lost) fifteen caudal vertebrae (AMNH online)
(AMNH 469; lost) twelve caudal vertebrae (AMNH online)
(AMNH 496) tibia (466 mm) (Pickering, 1996)
(AMNH 507) premaxilla, maxilla, teeth (Chure, 2000a)
(AMNH 530) femur, tibia, fibula (AMNH online)
(AMNH 600) incomplete skull (810 mm) (Osborn, 1900)
(AMNH 622) five cervical fragments (AMNH online)
?(AMNH 623) including partial distal caudal vertebra (AMNH online)
?(AMNH 624) including manual phalanx I-1 (AMNH online)
(AMNH 666; intended holotype of Allosaurus "whitei") incomplete
skull (885 mm), partial mandible, hyoid, axis, third cervical vertebra, fourth
cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh
cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth
cervical vertebra, first dorsal vertebra, second dorsal vertebra, third dorsal
vertebra, fourth dorsal vertebra, fifth dorsal vertebra, sixth dorsal vertebra,
seventh dorsal vertebra, eighth dorsal vertebra, ninth dorsal vertebra, tenth
dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal vertebra, sacrum,
ilium, pubes, proximal ischia (Osborn, 1900)
(AMNH 680; = AMNH 630 of Carrano, 1998?) (9.7 m; 2.3 tons) fifth to seventh
dorsal vertebrae, second to fourth caudal vertebrae, ~sixth caudal vertebra,
ilium, pubes, ischia, femur (1.008 m), tibiae (856 mm), fibulae, astragali,
calcaneum, metatarsus (IV 342 mm), pedal phalanges (Chure, 2000a)
(AMNH 813) two anterior dorsal vertebrae, five posterior dorsal vertebrae, dorsal
ribs, sacrum, two proximal caudal vertebrae, ilium, pubes, ischium (Chure, Fiorillo
and Jacobsen, 2000)
(AMNH 839) lower limb element (AMNH online)
(AMNH 847) five or six incomplete caudal vertebrae (AMNH online)
(AMNH 851) mandible (Chure, 2000a)
(AMNH 865) teeth (AMNH online)
(AMNH 857) tibia (AMNH online)
(AMNH 858) teeth (AMNH online)
(AMNH 865) premaxilla (AMNH online)
(AMNH 960) premaxilla, maxilla, teeth (AMNH online)
(AMNH 5750) teeth, three vertebrae, metacarpal, tibia (667 mm), fibula, ungual,
fragments (Carrano, 1998)
(AMNH 5753; =4753 of Glut, 1997) partial skull (maxilla, jugal, quadratojugal,
quadrate, ectopterygoid, partial pterygoid, braincase), axis, third cervical
vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth cervical
vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical
vertebra, tenth cervical vertebra, first dorsal vertebra, second dorsal vertebra,
third dorsal vertebra, fourth dorsal vertebra, fifth dorsal vertebra, sixth
dorsal vertebra, seventh dorsal vertebra, eighth dorsal vertebra, ninth dorsal
vertebra, tenth dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal vertebra,
thirteenth dorsal vertebra, sacrum, mid caudal vertebra, chevron, scapula, coracoid,
furcula, ilium, pubes, ischium, femur (Matthew 1908)
(AMNH 6125) femur (868 mm), tibia (758 mm), fibula, metatarsals (III 355, IV
325 mm), two pedal phalanges, two pedal unguals (Carrano, 1998)
(AMNH 6128) seven phalanges (Pickering, 1996)
(AMNH 30798; Dracula) (6.4 m) skull, mandibles, incomplete skeleton missing
some dorsal vertebrae and all caudal vertebrae (Galiano and Albersdorfer, 2010)
(AMNH coll.) twelve teeth (Brown, 1935)
(MOR 693; Big Al) (1.5 tons) skull, sclerotic ring, mandibles, cervical series,
cervical ribs, dorsal series, dorsal ribs, gastralia, sacrum, first two caudal vertebrae,
first chevron, scapulae, coracoid, humerus (318 mm), radii, ulnae (254 mm), manus including
manual phalanx I-1 and metacarpal II (118 mm), ilia (640 mm), pubes, ischium,
femora (742 mm), tibiae (665 mm), fibulae, astragalus, pes including pedal ungual
II, metatarsal III (346 mm), pedal phalanx III-1, metatarsal IV (286 mm) and
metatarsal V (Breithaupt, 1996)
(NAMAL coll.) (4 m; juvenile) (skull 360 mm) maxilla, prefrontal, postorbital,
jugals, quadratojugals, squamosals, quadrates, vomer, palatine, pterygoids,
partial braincase, dentaries, splenials, surangulars, prearticulars, articular,
hyoids, cervical vertebrae 3-10, dorsal vertebrae 1-13, ribs, gastralia, sacrum,
caudal vertebrae except some mid caudals, pectoral girdle, forelimbs, hindlimb,
skin impression (300^2 mm) (Pinegar, Loewen, Cloward, Hunter and Weege, 2003)
(NAMAL coll.) (8.8 m; adult) material including pubis (Pinegar, Loewen, Cloward,
Hunter and Weege, 2003)
(SCGM0177; Caesar) partial skeleton including incomplete skull and partial mandibles
(Tucker, 2009)
(SMA 0005; = SMA 005/02; Big Al 2) (7.6 m, adult) almost complete skeleton, skin (Evers et al., 2013)
about fifty-five teeth (Galiano and Albersdorfer, 2010)
(UMNH VPC 481) material including jugal (Carrano et al., 2012)
(USNM 544100) skull (Paulina-Carabajal and Coria, 2015)
(YPM 1879; holotype of Camptonotus amplus) distal tarsal III, distal
tarsal IV, partial metatarsal I, phalanx I-1 (66 mm), metatarsal II (305 mm),
phalanx II-1 (117 mm), phalanx II-2 (92 mm), pedal ungual II (103 mm), metatarsal
III (345 mm), phalanx III-1 (124 mm), phalanx III-2 (87 mm), phalanx III-3 (60
mm), metatarsal IV (310 mm), phalanx IV-1 (84 mm), phalanx IV-2 (69 mm), phalanx
IV-3 (44 mm), phalanx IV-4 (32 mm) (Marsh, 1879)
?...(YPM 1892) maxillary fragments, postorbital, partial quadrate (220 mm),
incomplete supraoccipital, pterygoid fragment, jaw fragments, cranial fragments,
eight teeth, partial posterior cervical or anterior dorsal vertebra, fused first
and second sacral centra (Galton, Carpenter and Dalman, 2015)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Colorado, US
(AMNH 5778) tooth (AMNH online)
(CM 2045) femur (McIntosh, 1981)
(LACM 135661) tooth fragment (LACM online)
(USNM 8257) manual ungual II (Gilmore, 1920)
?(YPM 4845) sacrum (YPM online)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Montana, US
?(YPM 16975) three centra (Douglass, 1909)
tibia, fibula, three metatarsals, phalanx (Cooley and Schmidt, 1998)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, New Mexico, US (Exter quarry)
material (Foster, 2003)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, South Dakota, US (Wonderland Quarry)
teeth, distal metatarsal IV (Forster, 1996)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Utah, US (Hanksville-Burpee quarry)
(BYU 9466) incomplete specimen including partial skull (Smith et al., 1999)
?(CM 21736) scapulocoracoid (McIntosh, 1981)
(DNM C4) femur (880 mm) (Madsen, 1976)
(DNM D4) femur (905 mm) (Madsen, 1976)
(DNM 4741) ilium (Meyers and Hoops, 1993)
(DNM 4818) humerus (Meyers and Hoops, 1993)
(DNM 4822) (juvenile) ulna (Meyers and Hoops, 1993)
(LACM 154771) tooth (LACM online)
partial tibia (Gregory, 1938)
material (Mathews, Williams, Bonnan and Henderson, 2009)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Wyoming, US (Aurora Quarry 3 and 4, Cheryls' Blind,
Little Houston Quarry, Reed's Quarry 12, Quarry R, Quarry B, Quarry C, Quarry
D, Red Mountain, Something Interesting Quarry, Westphal Quarry)
(AMNH 704) dorsal vertebrae, few ribs, sacrum, nine caudal vertebrae, pelvis
(Chure, 2000a)
(AMNH 715) femur (AMNH online)
(AMNH 726) ischium (Pickering, 1996)
(AMNH 728) pubes (Chure, 2000a)
?(AMNH 736) proximal caudal centrum (Pickering, 1996)
?(AMNH 737) partial mid caudal vertebra (Pickering, 1996)
?(AMNH 780) mandible (AMNH online)
(AMNH 871) rib (AMNH online)
(AMNH 885) rib (AMNH online)
(AMNH 889) rib (AMNH online)
(AMNH 5752; in part) (Pickering, 1996)
?(AMNH 30138) phalanx, fragment (AMNH online)
(CM 82) proximal caudal centrum (McIntosh, 1981)
(CM 36037) caudal vertebra (McIntosh, 1981)
(KU coll.) (juvenile) metatarsals (Bader, 2003)
(SDSM 25248) (adult) cranial elements including, premaxilla, maxilla/dentary
fragment, jaw fragments, two teeth (Foster and Martin, 1994)
(SDSM 30510) (juvenile) partial basicranium, tooth, three cervical vertebrae,
dorsal vertebra, several ribs, two sacral vertebrae, four caudal vertebrae,
manual phalanx, manual ungual, ilia (200 mm), ischium, femur (274 mm), tibia
(295 mm), metatarsal IV (148 mm), several pedal phalanges, pedal unguals (Foster
and Chure, 1999)
?(SDSM coll.) several teeth (Foster and Martin, 1994)
(USNM 8367) atlas, axis (89 mm), third cervical vertebra (105 mm), fourth cervical
vertebra (106 mm), fifth cervical vertebra (111 mm), sixth cervical vertebra
(121 mm), seventh cervical vertebra (125 mm), eighth cervical vertebra (120
mm), ninth cervical vertebra (122 mm), tenth cervical vertebra, partial cervical
ribs 2-10, first dorsal vertebra, second dorsal vertebra (89 mm), third dorsal
vertebra (78 mm), fifth dorsal vertebra (85 mm), sixth dorsal vertebra (80 mm),
seventh dorsal vertebra (88 mm), eighth dorsal vertebra (87 mm), ninth dorsal
vertebra (94 mm), tenth dorsal vertebra (96 mm), eleventh dorsal vertebra (99
mm), twelfth dorsal vertebra, thirteenth dorsal vertebra (102 mm), eleven dorsal
ribs, gastralia, fourth sacral vertebra (152 mm), fifth sacral vertebra (132
mm), first caudal vertebra (121 mm), first chevron, second caudal vertebra (123
mm), second chevron, third caudal vertebra (125 mm), third chevron, fourth caudal
vertebra (120 mm), fourth chevron (252 mm), fifth caudal vertebra (118 mm),
fifth chevron, sixth caudal vertebra (120 mm), seventh caudal vertebra (120
mm), seventh chevron, mid caudal vertebra (144 mm), partial ilium, pubes (740
mm), ischia (650 mm) (Gilmore, 1920)
(USNM 8405) (sacrum 575 mm), first sacral vertebra (120 mm), second sacral vertebra
(96 mm), third sacral vertebra (107 mm), fourth sacral vertebra (125 mm), fifth
sacral vertebra (118 mm), manual phalanges, metatarsal, pedal phalanges (Gilmore,
1920)
(UW coll.) several dorsal vertebrae, pelvic elements, hindlimb, partial pes
(Hunter and Breithaupt, 2005)
(UW coll?) teeth (Levitt, 2007)
(WDC ATWA-5000) lateral tooth (17.1x8.9x6.9 mm) (Diepenbrock, 2020)
(WDC-DMP-02) partial skeleton including several cervical vertebrae, incomplete
dorsal column, partial caudal series, appendicular elements including humerus
(370 mm) (Birkemeier, 2011)
(WDC SI-686) lateral tooth (28.4x14.3x6.2 mm) (Diepenbrock, 2020)
(WDC SI-870) premaxillary tooth (19.28.1x9.5 mm) (Diepenbrock, 2020)
(WDC SI-875) premaxillary tooth (27.2xx9.8x11.4 mm) (Diepenbrock, 2020)
(WDC SI-893) premaxillary tooth (33.5x12.8x15 mm) (Diepenbrock, 2020)
(WDC SI-910) lateral tooth (21.6x11.7x7.8 mm) (Diepenbrock, 2020)
(WDC SI-1111) premaxillary tooth (29.4x10.4x11.3 mm) (Diepenbrock, 2020)
(WDC SI-1235) posterior premaxillary tooth (18.5x9.3x7.7 mm) (Diepenbrock, 2020)
(WDC SI-1167) lateral tooth (29.2x12.9x5.9 mm) (Diepenbrock, 2020)
(WDC SI-No#) premaxillary tooth (21.4x8x9.1 mm) (Diepenbrock, 2020)
(WDC SI-Unk2) premaxillary tooth (12.8x4.5x8.5 mm) (Diepenbrock, 2020)
(WDC-TY A) (~8 m subadult) premaxilla, maxilla, dentary, dorsal vertebrae, caudal
vertebra, scapulae, phalanx I-1, manual ungual I, fibula, phalanx II-1, pedal
elements (Goodchild Drake, 2004)
(WDIS 011) skeleton including quadrate (Bakker, 2000)
(WDIS 091 or WDIS 911) partial quadrate (Bakker, 2000)
(WDIS 536) maxillary tooth (58 mm) (Bakker, 2000)
(YPM 1894) vertebra, humerus, pubes, femur, tibia (Ostrom, 1969)
(YPM 1895; lost?) pubis (YPM online)
(YPM 2792) tooth (YPM online)
(YPM 4679) two metatarsals (YPM online)
(YPM 4680) tooth (YPM online)
(YPM 4840) vertebrae, ilium (Ostrom and McIntosh, 1966)
(YPM 4841) dorsal centrum (YPM online)
(YPM 6222) pubis (YPM online)
(YPM 6223) pubis (Chure, 2000a)
(YPM 6224) pubis (Chure, 2000a)
(YPM 6225) ischium (YPM online)
(YPM 6226) (YPM online)
(YPM 40914; = YPM 1876) metacarpal (YPM online)
(YPM 46147) manual ungual (YPM online)
(YPM 46148) tooth (YPM online)
(YPM 46149) tooth (YPM online)
(YPM 54357) three lateral teeth (YPM online)
(YPM 54361) anterior tooth (YPM online)
(YPM 56468) caudal vertebra (YPM online)
(YPM 56469) caudal vertebra (YPM online)
(YPM 56470) caudal vertebra (YPM online)
(YPM 56471) phalanx (YPM online)
(YPM 56472) caudal vertebra (YPM online)
(YPM 56473) caudal vertebra (YPM online)
(YPM 56474) caudal vertebra (YPM online)
(YPM 56475) caudal vertebra (YPM online)
(YPM 56476) caudal vertebra (YPM online)
(YPM 56477) caudal vertebra (YPM online)
(YPM 56478) caudal vertebra (YPM online)
(YPM 56479) caudal vertebra (YPM online)
(YPM 56480) vertebra (YPM online)
(YPM 56481) caudal vertebra (YPM online)
(YPM 56482) caudal vertebra (YPM online)
(YPM 56483) caudal vertebra (YPM online)
(YPM 56484) caudal vertebra (YPM online)
(YPM 56485) caudal vertebra (YPM online)
(YPM 56486) caudal vertebra (YPM online)
(YPM 56487) caudal vertebra (YPM online)
(YPM 56488) partial distal caudal vertebra (YPM online)
(YPM 56489) dorsal vertebra (YPM online)
(YPM 56490) dorsal vertebra (YPM online)
(YPM 56491) caudal vertebra (YPM online)
(YPM 56492) vertebra (YPM online)
(YPM 56493) vertebra (YPM online)
(YPM 56494) caudal vertebra (YPM online)
(YPM 56495) ungual (YPM online)
(YPM 56496) caudal vertebra (YPM online)
(YPM 56497) caudal vertebra (YPM online)
(YPM 56498) caudal vertebra (YPM online)
(YPM 58258) (juvenile) proximal femur, proximal tibia (Dalman, 2014b)
(YPM 58259) (juvenile) sixth dorsal centrum (60 mm) (Dalman, 2014b)
(YPM 58260) (juvenile) partial lacrimal, two angular fragments, distal radius,
distal fibula, six postcranial fragments, five fragments (Dalman, 2014b)
(YPM 58261) partial ulna (Dalman, 2014b)
(YPM 58273) distal caudal centrum (Dalman, 2014b)
(YPM 58275) (juvenile) distal metatarsal III (Dalman, 2014b)
(YPM 58276) (juvenile) incomplete manual phalanx III-1 (Dalman, 2014b)
(YPM 58277) (juvenile) incomplete manual ungual III (Dalman, 2014b)
(YPM 58278) (juvenile) tooth, vertebra (YPM online)
incomplete skeleton including ilium (811 mm) and pubis (Chure and Fiorillo,
1997)
(embryos) bones including two premaxillae, eggshells, nest (Carrano, Mateus
and Mitchell, 2013)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, US
('BYU 5524'; Scheetz pers. comm. 2014 notes no BYU specimen corresponds to this)
ischium (Perez-Moreno et al., 1999)
('CM 10936'; Peratherium specimen in CM online catalog) humerus (Smith
et al., 1999)
(DMNS EPV21968) tooth (Kane, 2020)
(DMNS coll.) nineteen teeth (Kane, 2020)
(NMMNH 26086) tooth (43x21.3x? mm) (Heckert et al., 2003)
(NMMNH 26087) tooth (31.8x18.1x? mm) (Heckert et al., 2003)
(SMM 66-42-1) humerus (Smith et al., 1999)
(UMEM coll. = 'UMP' coll.) humerus (Smith et al., 1999)
(UNL 50038) humerus (Smith et al., 1999)
(UNL 50039) humerus (Smith et al., 1999)
(UNL uncatalogued) dentary (Smith et al., 1999)
(WDIS coll.; Rip Van Al) specimen including two dorsal vertebrae (Rothschild
and Tanke, 2005)
('YPM 1333'; Clidastes specimen in YPM online catalogue) premaxilla (Smith
et al., 2005)
('YPM 6293'; Orthacanthus specimen in YPM online catalogue) (Chure, 2000a)
incomplete postcranial skeleton (Paton, 1975)
numerous specimens from numerous localities (quarry list in Turner and Peterson,
1999)
Kimmeridgian, Late Jurassic
Porto Novo Member of Lourinha Formation, Portugal
(ML 415; holotype of Allosaurus europaeus) posterior
skull, sclerotic ring, posterior mandible, fourth to sixth cervical vertebrae,
fourth to sixth cervical ribs (Mateus et al., 2006)
Kimmeridgian, Late Jurassic
Camadas de Alcobaca Formation, Portugal
(IPFUB Gui Th 4) (juvenile) maxilla (23 mm) (Rauhut and Fechner, 2005)
(MNHNUL/AND.001) (~6.5 m) partial frontal, partial quadrate, teeth, ~eleventh
or twelfth dorsal centrum, ~thirteenth or fourteenth dorsal vertebra, two mid
dorsal ribs, gastralial fragments, first sacral centrum fused to partial second
sacral centrum, incomplete ~fourth caudal vertebra, partial mid caudal vertebra,
five distal caudal vertebrae, chevrons, ilial fragment, pubes (one incomplete),
incomplete ischium (~410 mm), incomplete femora, tibiae (one incomplete), fibulae,
astragalus, calcaneum, metatarsal II, metatarsal III, metatarsal IV, pedal phalanges,
pedal unguals (Perez-Moreno et al., 1999)
?....(MNHNUL/AND coll.) nasal fragment, lacrimals, quadratojugal, quadrate,
frontal, braincase, dentary, posterior mandible, cervical vertebrae, cervical
ribs, dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, chevrons,
coracoid, manual phalanges, ilium, pedal phalanges (Malafaia et al., 2010)
(MNHNUL/AND coll.) ilium (Malafaia et al., 2010)
Comments- Note that most of the above material has not been described,
and much is probably not distinguishable from Saurophaganax based on
what little we know of the latter genus. Indeed, as Allosaurus is the
most common Morrison theropod, it is likely material is often ascribed to it
without good reason, so some specimens may belong to Ceratosaurus, Marshosaurus,
Torvosaurus, Stokesosaurus, etc.. There is abundant material from
numerous quarries, not all of which are guaranteed to be listed above, though
all states and members are covered. For details on these, see Turner and Patterson
(1999) and Foster (2003).
The types- The holotype was discovered in 1877, and is from the Tithonian
Brushy Basin Member (Felch Quarry 1) of the Morrison Formation in Colorado.
It consists merely of an incomplete tooth, incomplete cervical or anterior dorsal
centrum, two post-cervical centra (posterior dorsal and proximal caudal in Marsh
1877; posterior dorsal and posterior sacral in Gilmore, 1920; dorsal and posterior
dorsal in Chure, 2000; caudals in Loewen and Chure, 2010), two rib fragments,
a humeral fragment and pedal phalanx III-1. Marsh diagnosed this by amphicoelous
(postcervical) centra transversely and ventrally reduced in size in the middle
and slender "feet bones" (pedal phalanx III-1). These characters are
plesiomorphic for theropods. Although partially figured by Gilmore, Madsen (1976)
and Glut (1997), the material has only been described by Chure. Prior authors
have viewed the holotype in different ways- Gilmore felt it was "hardly
more diagnostic" than Antrodemus, Madsen thought the humerus was
specifically diagnostic, and Chure admitted it could not be distinguished from jimmadseni but depended on his topotype. The tooth, unprepared presacral
centrum, rib fragments and pedal phalanx cannot be identified past at least
Carnosauria based on published information. One incomplete dorsal centrum resembles
the fourth dorsal (fifth in Madsen, 1976) most based on the centrum elongation
and shallow ventral concavity, though its broad ventral surface is first found
on dorsal six of DINO 11541. It can be distinguished from carcharodontosaurids
based on the lack of pleurocoels and from metriacanthosaurids based on its greater
width. If this is a fourth dorsal, its lack of pleurocoels would also distinguish
it from Saurophaganax, which seems to have them extend through dorsal
five. The longer centrum seems to be a fifth sacral based on its width (minimum
width 79% of length), in agreement with Gilmore's identification as a posterior
sacral but contrasting with Chure's identification as a posterior dorsal. It
is broader than metriacanthosaurids and Neovenator, and lacks pleurocoels
present in Acrocanthosaurus. Chure states Allosaurus has shallow
central fossae in its sacrals (visible in USNM 4734- Gilmore, 1920) while Saurophaganax
lacks them in sacral five, and these are present in the Allosaurus holotype.
Note that while Loewen and Chure (2010) have most recently identified both of
these centra as caudals without comment, Allosaurus has a narrow ventral
margin on its caudal vertebrae whereas these are broad. Madsen concentrated
on the humeral fragment to support the holotype's diagnostic nature, referencing
the position of the m. humeroralialis scar and nutrient foramen. Unfortunately,
carnosaur humeri are seldomly preserved and often not illustrated or described
in detail. Acrocanthosaurus and Mapusaurus both have a more posteriorly
placed scar and the former at least has a more anteriorly placed foramen. Acrocanthosaurus
also has a more abruptly expanded deltopectoral crest. However, Saurophaganax
has an anteriorly placed scar as in the holotype, though the position of its
foramen cannot be determined from publications. This suggests the humerus is
at best identifiable to Allosauridae based on published information, and not
a particular genus. This review suggests that the holotype might be referred
to Allosaurus based on the lack of pleurocoels in dorsal ~4 and the the
lateral fossae in sacral 5, but these are based on estimated positions in the
holotype's and Saurophaganax's material.
Madsen (1976) tried to designate DINO 2560 (then UUVP 6000) as a neotype, but
this is not allowed when the holotype is still extant (ICZN Article 75.3.4).
The nearly complete specimen USNM 4734 was designated the topotype by Chure
(2000) as it comes from the same quarry as the holotype, but the ICZN does not
recognize topotypes as official and the publication is a thesis. Most recently,
Paul and Carpenter (2010) have petitioned the ICZN to make USNM 4734 the neotype
of Allosaurus fragilis, which was still being voted on as of 1-2-2015.
There are a number of issues with the petition however. Paul and Carpenter fail
to list a centrum, humeral fragment and the rib fragments (noted by Demirjian,
2010 and Loewen and Chure, 2010) and continue to identify both free centra as
dorsals, which is seemingly incorrect for at least one as noted above. They
simply state the holotype elements "are inadequate to diagnose a genus,
much less a species, YPM 1930 can only be identified to family level" without
any detailed examination or reference to such an examination. As detailed above,
the holotype may share characters with Allosaurus to the exclusion of
Saurophaganax (the only other commonly accepted allosaurid genus), a
genus the authors never mention. Paul and Carpenter declare numerous species
to be indeterminate without providing evidence, some of which are also said
to be probably distinct from the holotype's species, also without evidence.
Regardless of these deficiencies and the lack of any detailed published comparative
study of the holotype, USNM 4734 will probably be declared the neotype based
on past ICZN action.
USNM 4734 was discovered in 1883 from the locality of the holotype. This was
first reported and illustrated by Marsh in 1884 (Gilmore, 1920), contrary to
Chure (2000) stating parts were illustrated by Marsh in 1879, prior to its actual
discovery (the pubis illustrated by Marsh 1879 is the YPM specimen shown in
Gilmore's plate 11, while the ischium's identity remains uncertain). Only the
pelvis and hindlimb are from USNM 4734 however, the humerus being YPM 1894,
and the rest based on Ceratosaurus and Megalosaurus. Gilmore (1915)
described the actual pectoral girdle and forelimb of USNM 4734 and later (1920)
described much of the rest of the specimen, though note most of the axial description
is of USNM 8367. Chure notes the right dentary on the specimen is actually from
USNM 8335 while Loewen and Chure (2010) state the premaxilla is too large and
thus from another individual. The latter authors also provide a materials list
for USNM 4734, though because Gilmore says not all carpals were found for each
side and leaves some measurements unknown for the left manus, their description
of the hindlimbs as complete may be incorrect. USNM 4734 can be assigned to
Allosaurus instead of Saurophaganax based on having atlantal prezygapophyses,
a semilunate atlantal intercentrum, dorsal pleurocoels limited to the first
two centra, posterior dorsals lacking a paraspinal lamina, and the tibia having
an astragalar buttress and a poorly projected medial malleolus.
Short-snouted fragilis vs. long-snouted atrox?- In the 1980s and 1990s it was common (e.g. Paul, 1988; Britt, 1991) to recognize two types of normal-sized
Morrison Allosaurus- one with a shorter snout and pointed lacrimal horns
(usually called A. fragilis) and another with a long snout and rounded
lacrimal horns (usually called A. atrox, and sometimes separated as Creosaurus).
While A. fragilis sensu stricto has been based mainly on the proposed
A. fragilis neotype, the 'A. atrox' morphology has not been based
on the Creosaurus atrox holotype but instead probably originates in Osborn
(1903) describing the elongate skulls AMNH 600 and 666 as Creosaurus.
Thus, actual discussion of the Creosaurus holotype is separated in its
own entry here, while the long-snouted morphotype will be called 'A. atrox'
for this section. Regarding the supposedly shorter skull of A. fragilis,
Smith (1998) first noted USNM 4734 "does not have any significant differences
from the articulated crania from the other sites at the individual element level
... Given the results presented here, a case could be made that this specimen
might be more correctly reconstructed as having a longer snout." Such a
case was made by Chure (2000), who notes that is the only short skull known,
and it was found disarticulated. When it was reconstructed by Gilmore (1920),
he had to "comprimise in regard to the exact articulation of the elements".
There are large plaster filled gaps in the specimen, the contact between the
maxilla, jugal and lacrimal is missing, the dentary is from another specimen,
the other mandible is plaster, the palate is fragmentary, and the postorbital
regions are distorted judging by their asymmetry. Chure notes the maxilla is
reconstructed too far posteriorly, as the lacrimal articulation of the dorsal
process is projecting into the antorbital fenestra. The angle between the maxillary
body and its dorsal process is similar to other Allosaurus specimens,
which wouldn't make sense if the snout were shorter. Similarily, the angle between
the anterior and ventral lacrimal processes is in the middle of the range Allosaurus
exhibits, with Cleveland-Lloyd 'A.atrox' specimens showing marked
variation. The nasal of USNM 4734 is broken and the anterior part moved dorsally
and rotated ventrally. The lacrimal horn shape shows many intermediates between
tall and triangular (USNM 4734) and low and rounded (DINO 2650), and there is
an example of a triangular lacrimal on a long skull (MOR 693). Contra Paul,
triangular lacrimals are known from the Cleveland-Lloyd quarry (eg. UUVP 40-581).
Though Paul claimed 'A. atrox' has a more robust neck, there is no difference
when cervical width/length ratios are compared. Similarily, though Paul claimed
'A. atrox' has a more robust forelimb, no difference was noted when humeral
circumference and length were quantitatively compared (circumference/length
ratio .45 in A. fragilis, .36-.49 in 'A. atrox'). Finally, both
supposed species are found in the same quarry, as evidenced by AMNH 600 (referred
to A. fragilis by Paul) and AMNH 666 (which he referred to 'A. atrox').
This is contrary to the stratigraphic distinction supported by Bakker and others.
In conclusion, there is no evidence for the fragilis/atrox dichotomy
advocated by Paul and Bakker, though it continues to be used in taxonomic arguments
(e.g. Paul and Carpenter, 2010).
Camptonotus amplus- Camptonotus amplus (Marsh, 1879) was
described based on an incomplete pes found in 1879. The diagnosis compared to
the type species C. dispar described in the same paper was a combination
of characters found in both species (reduced pedal digit I; fifth pedal digit
absent), those typical of Allosaurus (larger than Camptosaurus;
metatarsal I not contacting tarsus), one innaccurate character (pedal ungual
II proportionately longer- actually 34% of mtII length in amplus vs.
44% in dispar), one based on an incorrectly assigned element (pedal ungual
I more compressed transversely), and two that are too vague to evaluate (first
metatarsal "much curved"; pedal digits III and IV "large and
powerful"). Marsh (1885) later renamed the genus Camptosaurus, as
Camptonotus was preoccupied by gryllacridid cricket named by Uhler in
1864. Gilmore (1909) briefly commented on C. amplus in his Camptosaurus
monograph and provisionally referred the skull YPM 1887 to it because of its
large size (later named Theiophytalia kerri in 2007). Based on a personal
communication with Bakker, Galton and Powell (1980) referred amplus to
Allosaurus fragilis. This has been reconfirmed by McDonald (2011), and
the specimen finally received a modern description in Galton et al. (2015).
These authors confirmed the material belongs to Allosauroidea, except for the
pedal ungual I which they identify as sauropod and probably Camarasaurus,
recataloging it as YPM 58526. Additionally, they note Allosaurus specimen
YPM 1892 was found in the same quarry and is of comparable size, so probably
belongs to the same individual. Galton et al. provisionally create the new combination
Allosaurus? amplus, stating it differs from A. fragilis and Saurophaganax
in having a longer non-tapered portion of metatarsal I, and differs from A.
fragilis but not Saurophaganax in having a broad distal metatarsal
IV articular surface. They also posit the rather complete yet mostly undescribed
specimen AMNH 5753 may be referrable to A? amplus as it shares the broad
distal metatarsal IV.
The condition of metatarsal I is different from Madsen's composite UUVP figure,
DINO 11541 and 22 UMNH specimens mentioned by Galton et al., but basically every
well described Allosaurus specimen differs from others in some characters.
Similarly, narrow distal metatarsal IVs are reported in Madsen's composite and
DINO 11541, while wide ones are present in AMNH 5753, USNM 57589, YPM 1879 and
Saurophaganax specimen OMNH 1936. There is thus no established pattern
that width relates to taxonomy, as no two specimens referred to the same species
based on other characters have been shown to have the same condition. There
are also examples from each member of the Morrison with each morphology, so
the distinction is not stratigraphic and is here viewed as individual variation.
However, the distal end of metatarsal IV is triangular in amplus as in
Allosaurus (including USNM 57589; AMNH 5753's are unfigured) but not
the only figured specimen of Saurophaganax. Because of this, amplus
is here synonymized with Allosaurus fragilis pending defensible subdivision
of that species.
Interestingly, amplus is the first officially named allosaurid
from the Salt Wash Member, so if Loewen's (2009a) studies are correct that each member
has its own species, the lower member's species would be Allosaurus amplus
instead of A. jimmadseni (unless amplus is specifically indeterminate).
Thus it seems important to compare to jimmadseni's pes. Unfortunately,
the only pedal character identified by Chure (2000a) as differing between jimmadseni
and fragilis is "proximal corner of pedal phalanx III-2 projected
further laterally", which doesn't seem to actually be true. There are also
no characters preserved and published in YPM 1892 which can be compared to Chure's
distinctions between A. fragilis and jimmadseni. The metatarsi
are quite dissimilar as far as allosaurid specimens go, with amplus far
more robust, having a distolateral slope to the proximal surface vs. a distomedial
one, distally more bowed metatarsals III and IV which narrow and close the space
between them, a less ginglymoid metatarsal III, proximally broader metatarsal
II, posteriorly convex proximal outline of metatarsal IV, shallower posterior
intercondylar groove of metatarsal II, side edges of metatarsal III's distal
outline that are slanted ventromedially instead of ventrolaterally, a more convex
dorsolateral edge to the distal outline of metatarsal IV, plus the metatarsal
I and distal metatarsal IV distinctions noted by Galton et al.. Some of these
differences may be ontogenetic as amplus is 53% larger than jimmadseni,
but these specimens do nothing to support Salt Wash allosaurids being a coherent
species separate from A. fragilis.
Labrosaurus ferox- Labrosaurus ferox (Marsh, 1884) is based on
a pathological dentary (USNM 2315) from the Kimmeridgian Brushy Basin Member
(Felch Quarry 1) of the Morrison Formation in Colorado, found in 1883. It was
initially diagnosed by the toothless anterior (pathological), ventrally extended
posterior edge (also found in A. fragilis- MOR 693 in Chure, 2000) and
triangular teeth (unpreserved except for replacement tips). Hay (1908) recognized
that Labrosaurus ferox could not be compared to the genus' type species
lucaris, though he viewed the pathological morphology as natural so separated
it from Allosaurus. Gilmore (1920) was the first to suggest pathology
for the concave edentulous section of the dentary. Bakker (2000) suggested the
dentary "fits well into the anterior end of the post-dentary bones of USNM
4734 and certainly belongs to the same species if not the same individual",
and is from the same quarry. As such, this could be an objective junior synonym
of Allosaurus fragilis if USNM 4734 becomes the neotype, though it it
incomparable to Saurophaganax.
"Madsenius"- This name was orginally reported in a children's
book (Lambert, 1990) as "a proposed new allosaurid theropod to be formally
named and described." Olshevsky (1991) listed it under Allosauridae as
a taxon "to be described from the Morrison Formation by R. T. Bakker; based
on distinctive skull material and other remains previously referred to Allosaurus
and Creosaurus." Olshevsky (2004 online) wrote "trux"
"was to have been Bakker's original type species epithet for the as-yet-unpublished
genus Madsenius. According to him, it fits Madsen as appropriately as it fits
Madsenius" [etymology- Latin trux means "fierce, rough, savage, wild"].
Williams (2004 online) replied "This is essentially the version I heard too ~ "trux" = truculent."
While nothing else unique has been written regarding "Madsenius",
I believe clues in the literature point to its probable identity. Since at least
1988, Bakker has proposed two kinds of Morrison allosaur, the classic short-snouted
fragilis vs. long-snouted 'atrox' dichotomy. Bakker (2000) cited
the latter species as "The creosaur-type allosaurid (unfortunately, the
type of Creosaurus MARSH is, by itself, indeterminate): Dinosaur National
Monument skeleton University of Utah UUVP 6000 ..." as opposed to "True
Allosaurus MARSH: specimens from the type locality of Allosaurus fragilis
MARSH - the skeleton United States National Museum USNM 4734 ..." In that
paper, he stated "These two types of skulls are easy to tell apart from
the quadrate, lower temporal fenestra, and depth of the mandible; however, I
find it impossible to separate the two taxa from isolated snout bones or post-crania."
This matches the "distinctive skull material" noted by Olshevsky,
and the 'long-snouted' skulls have been referred to both Allosaurus and
Creosaurus by different workers, also matching Olshevsky's comment. Furthermore,
UUVP 6000 (later recatalogued as DINO 2560) was the basis of Madsen's (1976)
classic Allosaurus monograph and "Madsenius" clearly refers
to Madsen. Putting everything together, I think it's apparent "Madsenius"
was to be Bakker's name for creosaur-type allosaurs when he realized the Creosaurus
holotype couldn't be assigned to either variety. UUVP 6000 was probably supposed
to be the holotype.
If we accept this explanation, Bakker's characters supporting "Madsenius"
can be evaluated. Bakker states the ventral quadrate angles posteriorly in DINO
2560, forming a deeply concave posterior edge to the element unlike A. fragilis
AMNH 600. The posterior angle formed by the dorsal and ventral quadrate edges
is 24 degrees in AMNH 600 compared to 30 degrees in the UUVP coll. quadrate
illustrated by Madsen and 27 degrees in his cranial reconstruction of DINO 2560.
Bakker's DINO 2560 illustration shows an unprecedented angle of 52 degrees.
Angles in other specimens are 18 (AMNH 30798), ~44 (BYU 571/8901), 30 (DINO
11541), 43 (MOR 693), 15 (SMA 005/02) and 34 degrees (USNM 4734). For the laterotemporal
fenestra, Bakker states the restriction caused by the ventral squamosal process
is greater in A. fragilis AMNH 600 (least anteroposterior length of fenestra
15% of dorsoventral height) than DINO 2560 (26% in Madsen's reconstruction,
28% in Bakker's). Measurements in other specimens are ~25% (AMNH 666), 31% (AMNH
30798), <24% (BYU 571/8901), 38% (DINO 11541), 20% (ML 415), 40% (MOR 693),
38% (SMA 005/02) and 16% (USNM 4734). Mandibular depth is 19% of length in Bakker's
DINO 2560 (similar to 19% in Madsen's reconstruction) and 24% in his A. fragilis
illustration, though the latter is a composite between AMNH 666 (which has only
a partial surangular) and 5753 (which does not include mandibular elements).
So even this minor 5% difference cannot be determined. Values in other specimens
are ~19% (AMNH 30798), ~21% (BYU 571/8901), 17% (DINO 11541), 19% (MOR 693)
and 20% (SMA 005/02). The above comparison suggests mandibular depth is fairly
constant in known allosaurids, though quadrate angling and laterotemporal fenestra
proportions vary widely. Yet importantly, the latter conditions both exhibit
intermediates instead of two distinct clusters, and do not covary- AMNH 30798
and SMA 00/02 have low quadrate angles but wide laterotemporal fenestrae, while
USNM 4734 has a high angle but restricted fenestra. Note neither of these conditions
vary with stratigraphy either, and indeed the A. fragilis types and DINO
2560 are both from the Brushy Basin Member. Nor does it vary geographically,
with Wyoming specimens encompassing almost the entirity of the variation. Based
on this study then, "Madsenius" can be considered a synonym of Allosaurus
fragilis and another example of Bakker's notorious splitting.
Allosaurus "whitei"- This is based on a skull
with hyoid, presacral column, sacrum and pelvis (AMNH 666) discovered in 1901,
first mentioned in Osborn (1903) as Creosaurus. These were recovered
from the Kimmeridgian Salt Wash Member (Bone Cabin Quarry) of the Morrison Formation
in Wyoming. It was diagnosed by the same characters Paul (1988) used to distinguish
'A. atrox' (low, long skull; bluntly rounded lacrimal horn; slender limb
elements), so is invalid for the same reasons. Only the skull has ever been
illustrated (in Osborn, 1906 and 1912) and was only briefly described. Pickering
initially listed the name in a 1995 publication which he agrees only established
a nomen nudum, and later used it in more detail in a 1996 publication which
he sees as valid. Chure (2000a) rejected the validity of the name, as Pickering
didn't follow ICZN Article 8 Recommendation 8A (Authors have a responsibility
to ensure that new scientific names, nomenclatural acts, and information likely
to affect nomenclature are made widely known), Article 8.1.1 (it must be issued
for the purpose of providing a public and permanent scientific record), Article
8.1.2 (it must be obtainable, when first issued, free of charge or by purchase),
and a recommendation from the third edition of the ICZN absent in the fourth
edition (conventional printing being preferred, though the new Article 8.4.1
expressly allows Pickering's method). As recommendations are optional, only
Articles 8.1.1 and 8.1.2 are necessary for the name to be valid. Article 8.1.1
is about intent, and while Chure is correct that Pickering's 1996 work is far
from a traditional scientific work, his online remarks indicate new names there
were meant to be public, permanent and scientifically valid. Regarding Article
8.1.2, Chure states it was sent to "some unknown number of personal friends
of the author", but Pickering (online, 2002) stated "well over 150
copies of my publication were distributed as a supplement to Mike Fredericks'
PREHISTORIC TIMES in 1996". The former would not qualify under 8.1.2, though
the latter would. As for AMNH 666, the meager published information is insufficient
to assign it to Allosaurus or Saurophaganax, though several characters
of the latter genus (postorbital lacks rugosity; mid cervicals with nearly vertical
postzygapophyses; mid dorsals have well developed infraprezygapophyseal lamina;
mid dorsals with vertically oriented infrapostzygapophyseal lamina; pleurocoels
present through fifth dorsal centrum) could be checked in the material. Pickering
also designated AMNH 5753 as a topotype, which the ICZN does not recognize but
is considered a specimen from the same locality as the holotype. This is the
mounted specimen standing low over Apatosaurus vertebrae (AMNH 222),
but Chure notes it was from Aurora Quarry 3 so cannot be a topotype. Pickering
(1996) also lists AMNH 275, 290, 324, 496, 600, 680, 704, 726, 728, 736, 737,
851, 5752, 6125, 6128, MCZ 3897 and possibly DINO 2560 as referred specimens.
If "whitei" turns out to be a valid name, and amplus is specifically
indeterminate, and Loewen's studies are correct that Salt Wash specimens aren't
A. fragilis, "whitei" would be the next valid name for that
species. However, note AMNH 704, 726, 728, 736, 737, 5752 and 5753 are from
quarries that have not been placed in the Salt Wash Member with certainty, and
that MCZ 3897 and DINO 2560 are from the Brushy Basin Member so would be A.
fragilis on stratigraphic grounds.
"Wyomingraptor"- This name was published in the column 'Dr.
Bob's Dinofacts' in response to a question from a reader (Anonymous, 1997).
The author (possibly Bakker himself) suggested it for a Tate Geological Museum
specimen currently labeled Allosaurus. From 1997 until 2006, the Tate
Museum included a "Wyomingraptor" section in its exhibits page, stating
Bakker has proposed that name for a new genus of allosaur found at Como Bluff
including a photographed forelimb. In the PaleoWorld episode "Killer Raptors"
(episode 7 of season 4) aired in 1997, Bakker claims the only theropod preserved
in Nail Quarry is "Wyomingraptor" (though note this is untrue, as
the "Brontoraptor" material was also found there). The material (three
adults and numerous juvenile to adult teeth) was mentioned in Bakker (1997)
where he simply calls them Allosaurus. Hartman (DML, 2000) wrote that
Bakker has been "attempting to erect a new genus of allosaur, which he
dubs "Wyomingraptor." He has been using this name for some time, but
recently has found a specimen he thinks is different enough from the type(s)
to warrant generic distinction." Given Bakker's notoriety as a splitter,
the Nail specimens are likely to just be Allosaurus fragilis in any case.
The photographed forelimb is similar to A. fragilis USNM 4734 except
for being more robust, at least in metacarpal I, phalanx II-1 and II-2. Indeed,
the robust first metacarpal is similar Torvosaurus, though the elongate
phalanx I-1, radius and ulna are not. The forelimb is stated to be a cast, so
it's not certain how much is based on real Nail fossils. It's possible some
elements were scaled incorrectly from other specimens or are complete fabrications.
It's also possible some material such as metacarpal I actually comes from the
"Brontoraptor" individuals and that the forelimb is a chimaera. Further
evaluation awaits description of the Nail material, which has yet to be distinguished
from Saurophaganax either.
A. jimmadseni- DINO 11541 was discovered on July 15 1990 in
the Kimmeridgian Salt Wash Member of the Morrison Formation in Utah (quarry
DNM 116). Chure (2000a) described it in his thesis as a new species, Allosaurus
"jimmadseni", first published by Glut (2003). Several more recent sources have used
A.
"jimmadseni" as valid (Snively et al., 2004; Malafaia et al., 2007;
Rauhut, 2011; Dalman, 2014a and b). None of these establish the species
as they lack an explicit statement of novelty (ICZN Article 16.1), but
it was officially named by Chure and Loewen (2020) who also described
the skull, mandible, hyoid and axis. They also refer several
other specimens to jimmadseni-
all BYU Dry Mesa Quarry material "including:
BYU 4861, 5164, 5268, 5292, 5583, 11936, 13621, 16942, 17106, 17281",
MOR 693, SDSM 30510, SMA 0005, UMNH VPC 481 and USNM 544100. This
seems to be a continuation of work done by Loewen (2004), Brandau et
al. (2008) and Loewen (2009a, b), who performed a "morphometric
analysis of over 570 bivariate comparisons for 1,300 specimens" of
skulls and hindlimbs and found two Allosaurus
species, stratigraphically separated.
Note that a division between members
isn't exactly the same as Chure's hypothesis of jimmadseni, as Chure considered AMNH 507, 600, 666, 680, 851 and 6125 to be
A. fragilis
despite being from the Salt Wash Member, while Chure and Loewen (2020)
referred "all allosaurid material from the Dry Mesa Quarry, CO curated
at BYU" to A. jimmadseni despite being from the Brushy Basin Member. Chure and Loewen explain the latter as "Allosaurus jimmadseni
occurs below the "clay change" of Turner & Peterson (1999), except
for at DMQ, which occurs only two m above the "clay change"." This
might therefore correspond to the boundary between Dinosaur Zones 2 and
3 of Turner and Peterson, but the Bone Cabin quarry that yielded supposed A. fragilis
AMNH 600 and 666 is ~100 meters below the clay change so would create
significant stratigraphic overlap. Pinegar et al. (2003)
mentioned two NAMAL specimens they referred to A. jimmadseni. Skin impressions found with the juvenile consist of
2-3mm wide scales. Galiano and Albersdorfer (2010) reported an additional A. jimmadseni
specimen from the Dana Quarry of Wyoming. Nicknamed Dracula, it is reportedly
~70% complete, missing the tail and some dorsals. The AMNH has acquired it as
AMNH 30798.
Chure (2000a) diagnosed jimmadseni
based on- row of maxillary neurovascular foramina below antorbital
fenestra in antorbital fossa (also used in official diagnosis; also in
AMNH 600; only one foramen in MOR 693); axial intercentrum rotated
dorsally (also used in official diagnosis; not in MOR 693); axial
intercentrum with flared rim (also used in official diagnosis; not in
MOR 693); accessory ossifications on the anterior and posterior edges
of proximal caudal neural spines (these are merely the interspinous
ligaments, which are partially ossified in A. fragilis as well); anteroposteriorly elongate
obturator notch in pubis delimited by triangular processes (a further ossification
of cartilaginous areas in most Allosaurus, with e.g. the distal triangular
process present but more poorly developed in the YPM specimen in plate 11 of
Gilmore 1920; Chure notes AMNH 813 seems to have an enclosed obturator foramen,
which would be another example); proximal corner of pedal phalanx III-2 projected
further laterally (untrue compared to AMNH 324 and Madsen's 1976 UUVP coll.).
He also lists numerous supposed differences from A. fragilis-
premaxillary narial fossa less well defined (difficult to evaluate);
large foramen between lacrimal and jugal (not maxilla, contra Chure's
text) at posteroventral corner of antorbital fossa (placed entirely
within lacrimal by Chure and Loewen 2020; not in MOR 693 or SMA 0005);
elongate medial fenestra to maxillary antrum (not verified in other jimmadseni
specimens); rounded lacrimal horn (distortion?; also in most
Cleveland-Lloyd specimens- Carpenter, 2010); lateral pneumatic recess
of lacrimal absent (untrue); lateral vertical ridge on lacrimal horn
(untrue based on Chure and Loewen 2020); lacrimal horn not rugose
(untrue based on Chure and Loewen 2020); straighter ventral margin of
jugal (also used in official diagnosis and in Loewen 2004 and 2009;
also in AMNH 600; curvature same in SMA 0005, UMNH 8976 and 9084,
within fragilis range in BYU 5122);
parietals fused posteriorly (untrue); parietals
taller than wide in posterior view (untrue); basipterygoid recess well marked
and invasive (variable in fragilis- Chure and Madsen, 1996a); myohyloid
foramen nearly enclosed ventrally (variable in fragilis- Madsen, 1976);
teardrop-shaped internal mandibular foramen in anteroventral prearticular (not verified in other jimmadseni specimens); nineteen
dentary teeth (18 in BYU 2028 and SMA 0005; 18 and 19 in MOR 693); odontoid process of axis tall
and narrow in anterior view (seemingly untrue based on plate); third cervical
neural spine slanted posteriorly (also in AMNH 5753 and USNM 8367;
not known to be absent in any fragilis specimen); fourth through sixth
neural spines more anteroposteriorly elongate (untrue of c6 in USNM 8367); cervical
pleurocoels change in size throughout the vertebral column (not known to be absent in any fragilis
specimen); fifth and sixth cervical epipophyses transversely compressed (true
in c4, 5 and 7 in Madsen's 1976 UUVP coll.); presacral eleven with parapophysis
fully on centrum (also in AMNH 5753 and Madsen's 1976 UUVP coll.; in USNM 4734
"the upper edge reaches the neurocentral suture", so this is not much
different); hypapophysis on presacral 11 instead of 12 (also in AMNH 5753; on
both in Madsen's 1976 UUVP coll. and USNM 4734); less pronounced notch between
acromion and anterior coracoid edge (barely apparent if at all compared to USNM
4734); distal scapula not as expanded dorsally (barely
apparent if at all compared to Madsen's 1976 UUVP coll.); coracoid extremely
thin anteriorly (not verified in other jimmadseni specimens); more gracile ulna (limb gracility shows variation in A.
fragilis- e.g. humerus in Smith's 1998 study showing diameters varying from
~40 to ~90 mm in ~330 mm long elements); longer olecranon process (only appears
to be present due to the bone's gracility); olecranon process transversely thin (not verified in other jimmadseni specimens);
medial ulnar cotyle lower (I'm unsure what direction this refers to); straighter
ulnar shaft (not in MOR 693); long axes of proximal and distal ulnar ends
offset by ~45 degrees instead of ~90 (not verified in other jimmadseni specimens); vertical ridge above acetabulum (also
in atrox);
pubic boot less tall and massive (not in SMA 0005); medial surface of
low femoral head angled strongly ventrally (while initially appearing
odd, this is due to a combination of flatter medial and dorsal edges
and an actual angular difference in 20 degrees from e.g. the YPM
specimen in plate 11 of Gilmore 1920 or Madsen's 1976 UUVP coll.;
similar variation in angle and medial flatness are seen in Megalosaurus bucklandii); strongly developed
mediodistal crest on femur (actually similar to some A. fragilis
such as UNMH VP 7884 and 12231); less curved cnemial crest (untrue of
the right tibia). Chure and Loewen also listed straight
posteroventral jugal ramus of maxilla where it articulates with jugal
(10 degrees difference between ventral edges of posterior portion and
rest of maxilla in DINO 11541; 5 degrees in AMNH 600; 14 in USNM 4734;
16 in AMNH 666; 16 in SMA 0005; 19 in MOR 693; 26 in DINO 2560);
laterodorsal margin of nasal pinched into low crest continuous from
premaxilla to lacrimal (probably "nasals of the two species are
distinct" from Loewen 2004; seems valid- present in DINO 11541, MOR 693
and SMA 0005; not in DINO 2560, USNM 4734, or supposedly BYU 9466, UMNH
VP 7748 and 9149); posterior portion of dorsal surface of nasal
cup-shaped, producing a median peak in region of nasofrontal contact
(possibly part of "nasals of the two species are distinct" in Loewen
2004; also in USNM 4734); relatively taller lacrimal horns (also in
USNM 4734); jugal with straight to slightly-curved outline in dorsal
view (also in Loewen 2009 as "caudal portion of the skull ...
substantially increasing in transverse breadth"; also in USNM 4374);
well-developed distinct antarticular (not known to be absent in any fragilis specimen).
Unfortunately, the degree of variation in Allosaurus specimens
is almost never mentioned by Chure (2000a) or Chure and Loewen (2020)
and very few specimens have been decently described or
illustrated. The large scale work of Loewen (2009a) remains
unpublished and only available as abstracts. Several of the
characters found in DINO 11541 seem to be the consequence of greater
ossification and most are comparable to intraspecific variation in
other theropods or specifically vary within specimens from each
stratigraphic level, with the best candidate for a valid character
being the narrow nasal ridges. Even the characters that made it
to the official diagnosis of Chure and Loewen are largely absent in A. jimmadseni
paratype MOR 683 they describe (row of foramina in maxillary antorbital
fossa; straight posteroventral maxillary edge; rotated and flared axial
intercentrum), present in proposed A. fragilis
neotype USNM 4734 (transversely concave posterodorsal nasal surface;
tall lacrimal horn; lateral margin of jugal straight in dorsal view),
or present in all Allosaurus
(well developed antarticular). Given the overlap in anatomical
features for all characters except one that can be evaluated in more
than one specimen in each stratigraphic layer, A. jimmadseni is considered a junior synonym of A. fragilis
pending publication of Loewen's (2009a) thesis and/or future
publications of Loewen and Chure. The latter mention "a
postcranial description and a revision of genus Allosaurus [that] will be the subject of a future publication" and "a subsequent publication [where] we will review all named species of Allosaurus from North America in support of our view that there are only two valid species of Allosaurus in North America, Allosaurus fragilis and Allosaurus jimmadseni."
Allosaurus "carnegeii"- This name was used in
a caption for the cover (on page 3) of the June 2003 issue of Discover Magazine,
advertising a story inside by Levin and Gurney. As Levin near certainly didn't
write the cover description, the usually cited authorship of "Levin, 2003"
is probably in error and should be attributed to Discover editors. It is no
doubt a typo, perhaps confusing the name with Diplodocus carnegii. The
cover photo is the skull and anterior cervicals of the mounted CM 11844 from
the Tithonian Brushy Basin Member of the Morrison Formation (Carnegie Quarry)
of Utah. As detailed by McIntosh (1981), this specimen was discovered in 1913
and the skull on the mount is a cast of DINO 2560 (then UUVP 6000). As the first
two cervicals seem to be at least partially sculpted (e.g. elongate axial neural
spine), there may not even be any real material in the cover photo. The name
is certainly a nomen nudum, lacking description (ICZN Article 13.1.1), an explicit
statement of novelty (ICZN 16.1) and explicit designation of a type (ICZN 16.4.1).
CM 11844 has never been described, but the mount has no paraspinal lamina, relatively
unexpanded chevrons and a highly divergent metatarsal IV, so is Allosaurus
instead of Saurophaganax if these areas were based on real material.
If Loewen's studies are correct, stratigraphy would place CM 11844 and DINO
2560 as A. fragilis.
Allosaurus europaeus- Perez-Moreno et al. (1999) described MNHNUL/AND.001
as Allosaurus fragilis, that was later described in detail by Malafaia
et al. (2007). Ortega et al. (2009) and Malafaia et al. (2010) mentioned and
illustrated more material later excavated from this specimen, and the latter
notes a larger ilium is preserved as well. Mateus et al. (2006) briefly described
a partial skull and cervicals as the new species Allosaurus europaeus,
referring MNHNUL/AND.001 based on geography although the specimens lack overlapping
elements. Malafaia et al. felt the supposed autapomorphies for A. europaeus
were primitive for allosauroids and/or subject to individual variation in A.
fragilis. My comparison confirms this. A. europaeus was diagnosed
by Mateus et al. based on the following characters- maxilla forked posteriorly
(also in AMNH 600); truncated ventroposterior process of maxilla (also in AMNH
600 and DINO 11541); nasal with two pneumatic foramina (anterior foramen twice the size of
the posterior) (posterior actually larger); rugose dorsal rim of nasal (also
in many specimens such as BYU 2028 and SMA 0005); lacrimal horn narrow in
lateral view (also in UMNH 7786 and 7785); lateral lamina of lacrimal subtle (also in AMNH 600); jugal participation
in antorbital fenestra (also in some A. fragilis- Bakker pers. comm.
to Carpenter and Currie, 2000; argued by Malafaia et al., 2009 and 2010 to be
a misinterpretation based on a crack instead of a suture); posteroventral projection
of jugal more than twice posterodorsal projection (also in BYU 571/8901, DINO
11541 and USNM 4734); ventral tip of postorbital reaches lower rim of orbit
(untrue, but similar extent in BYU 571/8901); anterior tip of quadratojugal
anterior to laterotemporal fenestra (also in AMNH 600); squamosal contacts quadratojugal
in sigmoidal suture; squamosal projects ventrally into laterotemporal fenestra
(true in all Allosaurus); occipital condyle above squamosal-quadratojugal
contact (also in USNM 4734); palatine contacts pterygoid dorsoposteriorly (true
generally in Allosaurus- e.g. DINO 11541 and MOR 693); large anterior
surangular foramen (also in AMNH 4734). Only the larger
posterior nasal foramen and sigmoidal squamosal-quadratojugal suture differ
from the sample of A. fragilis
skulls checked. Of these, the size and number of nasal foramina varies
between specimens (e.g. one foramen in DMNH 2419; two right and three
left in USNM 4734; three foramina in BYU 2028), as common for pneumatic
features. The squamosal-quadratojugal shape is also variable in A. fragilis,
so that it is unsurprising the sigmoid line of A. europaeus' type is
different from the other mutually different examples. While one might expect
European and American Allosaurus to be specifically distinct, this is
so far unjustified by morphology. I follow Malafaia et al. in synonymizing the
species.
Allosaurus lucasi- The material of this species (YPM 57589) was
discovered in 1953 in the Tithonian Brushy Basin Member of the Morrison Formation
in Colorado (McElmo Canyon). This was first mentioned in an SVP abstract (Dalman
et al., 2012) as a taxon "distinct from Allosaurus and Saurophaganax",
though phylogenetic analysis placed it "within Allosauroidea, more closely
related to Allosaurus, Fukuiraptor and Neovenator."
Dalman (2014) described it as a new species of Allosaurus, A. lucasi.
Contrary to Dalman et al., no CT scans or braincase are mentioned in the description.
Dalman also correctly identifies five premaxillary teeth, whereas Dalman et
al. state only four are present. Unfortunately, YPM 57589 is highly fragmentary
and only partially prepared, and Dalman's description is poorly written and
full of errors.
The first supposedly distinctive character (also mentioned by Dalman et al.)
is a short premaxilla, which Dalman claims is equal in length and height. However
figure 2A clearly shows the length/height ratio to be ~144%, which is actually
more elongate than A. fragilis exemplar USNM 4734 (138%), let alone specimens
with even shorter premaxillae (e.g. 85% in AMNH 600). Both Dalman and Dalman
et al. cite a short and deep maxilla as a second derived character, but only
the anterior third is preserved so this is completely unknowable. Note that
the labeled margin of the antorbital fossa (eaof in figure 4A) is actually above
the base of the ascending process and that little if any of the preserved section
was under the actual antorbital fossa. Thus when Dalman later states the antorbital
fossa was not as ventrally extensive in lucasi, he's confusing the portion
below the ascending process for the portion under the fossa, and the structure
is the same as in other Allosaurus specimens. Dalman lists reduced anterior
and posterior quadratojugal processes as diagnostic, but the posterior process
is equally short in e.g. AMNH 600, while the anterior process is broken. The
bone is only visible as two sagittal cross sections still encased in matrix,
and one block of matrix is missing the ventral edge present on the other block
so that the anterior process appears shallow and more tapered (figure 6D, also
used in plate 1), but the actual bone is deeper and less tapered (rqjr in figure
6C) so comparable to A. fragilis. Dalman et al. similarly list a robust
quadratojugal as diagnostic, but this is an illusion caused by the incorrectly
restored anterior process. Dalman states the ventral margins of the anterior
and posterior quadratojugal processes are on a straight line, but the angle
is 27 degrees, similar to the 32 degree measure in USNM 4734 and more than in
DINO 11541 (as Dalman himself notes later in the text). Dalman's last diagnostic
character is the supposedly posteriorly placed lateral tibial condyle, but this
is very similar to USNM 4734 (posterior edge of condyle at 87% of anteroposterior
length in lucasi vs. 86% in fragilis).
Dalman also lists numerous features supposedly different from A. fragilis
in the text. The anterior edge of the premaxilla is said to be at a slightly
lower angle compared to the ventral margin, but both edges are incompletely
preserved and the angle varies widely in other Allosaurus specimens.
The premaxillae are claimed to make a narrower symphysis by five degrees, but
this is not only within likely individual and/or taphonomic variation, it's
probably not possible to measure that exactly in lucasi due to the medial
side being "cemented to the conglomeratic matrix." Dalman claims the
subnarial foramen is absent, but given the "extensive weathering of the
lateral surface" which has removed traces of neurovascular foramina, the
subnarial foramen is probably unpreserved as well. The dorsal quadratojugal
process is said to be at a right angle to the ventral process unlike a supposed
75 degree angle in A. fragilis, but some fragilis specimens have
a right angle too (e.g. DINO 2560). The same thing can be said of the supposedly
straight instead of anteriorly bent dorsal quadratojugal process, with the added
issue that the dorsal portion is broken off. The posterior end of the quadrate
is said to be slender and that of fragilis robust, but so much of the
bone has been eroded away (dorsal third, mandibular condyles, anterior portion
of pterygoid wing) that the posterior width is about the only dimension measurable,
so can't be related to any other dimension to measure robusticity. What little
is preserved of the bone shows no differences from fragilis. The pubic
peduncle is said to end "with a small protruding structure, which extends
throughout the entire anteroposterior length of the bone", supposedly unlike
A. fragilis, atrox or jimmadseni, but what this refers
to isn't obvious from the figure. Dalman says the pubic peduncle is shorter
and more robust than A. fragilis, but as only the distal end is preserved,
this is impossible to evaluate. Similarly, the supposedly more shallow posterior
concavity is impossible to know since the proximal half of the concavity is
unpreserved., and the supposedly more gracile and much thinner pubic shaft may
not be true since only two short shaft fragments are preserved. Metatarsal I
is claimed to be more massive, but its width is ~53% of metatarsal II's based
on figure 18, compared to 53% in USNM 4734. Dalman is correct in stating metatarsal
IV is less distally divergent than some A. fragilis (e.g. AMNH 324, DINO
11541), though this is similar to Saurophaganax and is also found in
some fragilis specimens (e.g. MOR 693).
Metatarsal IV's triangular distal end identifies this as Allosaurus instead
of Saurophaganax, and Dalman's characters are inaccurate, unknowable
or fall within the range of variation of A. fragilis. Thus it is referred
to that species here. If Loewen's studies are correct, YPM 57589's high stratigraphic
position would keep it referred to A. fragilis.
Dalman also refers YPM 57726, supposed dentary and splenial fragments, to A.
lucasi because they and the holotype "were found in close proximity
to each other and exhibit similar morphology." The supposed dentary fragment
is from a smaller individual than the holotype, not stated to share any characters
with it to the exclusion of other Allosaurus, and can't even be placed
compared to the holotype dentary fragment given the poor figures and description.
Although stated to be from the "distal" (= anterior?) portion of the
dentary, there is no obvious Meckelian groove and the tall and convex supposed
splenial articular surface is unlike any theropod. Dalman states the splenial
articular surface "is much deeper than it is in other species of Allosaurus",
but as no theropods have a similar structure there is more likely misidentification
to blame. The supposed splenial fragment has too little anterior taper to be
an Allosaurus splenial, though it does match the middle section of an
angular well. There's nothing suggesting these fragments come from the same
individual or even from Allosaurus.
Other specimens- Lisak (1980) described a snout in his thesis as Allosaurus cf. fragilis
that was redescribed by Smith and Lisak (2001). BYU 2028 is from the Tithonian
Brushy Basin Member of the Morrison Formation in Utah. The authors state it
is unlike most Allosaurus specimens in having a premaxilla which is taller
than long, more extensive pneumatization around the antorbital fossa, convex
ventral maxillary edge, better developed nasal ridge, and more concave ventral
dentary edge. They considered these features to be individual variation within
A. fragilis.
MOR 693 was discovered in 1991 in the Kimmeridgian Salt Wash Member of
the Morrison Formation inWyoming (Laws, 1993). The specimen is very
complete and has been nicknamed Big Al. Big Al has been the focus of
pathological research (e.g. Laws, 1996; Breithaupt, 2001; Hanna, 2002),
though no description of its morphology has been published. Another
rather complete specimen (SMA 005/02) was discovered by the same team
in the same quarry in 1996, and is nicknamed Big Al 2. It has been
mentioned only briefly in the literature, though Evers et al. (2013)
did comment on its pathologies in an abstract, later detailed by Foth
et al. (2015). Chure and Loewen (2020) state "SMA 0005 is
currently being described by scientists at
Ludwig-Maximilians-University in Munich, Germany" and will be donated to the University of Zurich (Hotz, 2020).
NMMNH P-26083 (Williamson and Chure, 1996) is a large specimen from the Tithonian
Brushy Basin Member of the Morrison Formation in New Mexico. Williamson and
Chure merely referred it to Allosauridae, not a particular genus. It is assigned
to Allosaurus instead of Saurophaganax because the femoral shaft
is straight in anterior view, the tibial facet for the astragalar ascending
process is well marked and the medial malleolus of the tibia is poorly expanded.
P-26083's large size invites comparison to Epanterias, but no preserved
elements are shared with the holotype. The strongly curved cnemial crest, twisted
tibial shaft, and flattened medial surface of the medial tibial condyle are
supposedly unique characters. However, twisted tibial shafts are almost universal
among theropods and A. fragilis exemplar USNM 4734 also has a concave
medial edge of its medial condyle. As the tibia is not figured in proximal view,
the cnemial crest's curvature cannot be evaluated.
Smith et al. (1999) described BYU 571/8901 from the Brushy Basin Member of the
Morrison Formation in Utah as an example of Allosaurus fragilis. It has
a perhaps pathologically fused ectopterygoid and pterygoid (right side only),
fused atlantal elements, less distinct coracoid tuber, and humerus with greater
torsion than other A. fragilis. Bybee and Smith (1999) note it has an
ossified sphenethmoid, very thin parasphenoid rostrum, posteriorly oriented
basisphenoid recess, coalesced cranial nerve foramina at base of occipital condyle,
and more horizontally oriented paroccipital processes than most A. fragilis
as well.
The referred eggs are Prismatoolithus coloradensis (Hirsch, 1994), as
shown by Carrano et al. (2013).
Not Allosaurus- The holotype of "Laelaps" trihedrodon
(Cope, 1877) is lost and was never illustrated and only briefly described, so
cannot be assigned to Allosaurus or another genus (Chure, 2000a; published
as Chure, 2001).
The holotype of Hypsirophus discurus (Cope, 1878) does not contain Allosaurus
elements, contra Glut (1997) and Paul and Carpenter (2010) (Chure, 2000a).
Rhodes and Currie's (2020) Table 1 lists "Allosaurus fragilis BYUVP 17550 Fused, partial pubes (apron and boot)", but Scheetz (pers. comm., 7-2022) reports "BYU 17550 is the Ceratosaurus pelvis (pelvis consisting of 5
sacral verts, both illia, fused pair of pubes and fused prox ends of
ischia) from the Brushy Basin Member of the Morrison, Dry Mesa Quarry,
western Colorado."
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