= 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
für Geologie und Paläontologie 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
für Geologie und Paläontologie Abhandlungen. 210(1), 41-83.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and
nomenclature of the major taxonomic categories of the carnivorous
Dinosauria (Theropoda). Journal of Vertebrate Paleontology. 19(1),
69-80.
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 Osmólska
(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. Most recently, Rauhut et al.
(2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it
as either a basal megalosaurid, a eustreptospondyline or sister to
Allosauria.
References- Lapparent and Zbyszewski, 1957. Les dinosauriens du
Portugal. Mémoires du Service géologique du Portugal. 2, 1-63.
Mateus,
1997. Eggs, nest and embryos of theropod dinosaur in Upper
Jurassic level of Lourinhã, 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
planètes, 325: 71-78.
Mateus, 1998. Lourinhanosaurus antunesi, a new Upper Jurassic
Allosauroid (Dinosauria: Theropoda) from Lourinhã (Portugal). Memórias
da Academia de Ciências de Lisboa. 37: 111-124.
Mateus, Mateus, Antunes, Mateus, Taquet, Ribeiro and Manuppella, 1998.
Upper Jurassic theropod dinosaur embryos from Lourinhã (Portugal).
Memórias da Academia de Ciências 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 Lourinhã (Portugal). Comptes
rendus de l'Académie des sciences - Série IIa - Sciences de la Terre et
des planètes. 332(10): 647-656.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle
découverte et révision systématique: Implications phylogénétiques et
paléobiogéographiques. 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 Osmólska. 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 Ricqlès, in preparation. The osteology of
the embryos of the theropod dinosaur Lourinhanosaurus antunesi
from Portugal.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod
dinosaur from the Callovian Balabansai Formation of Kyrgyzstan.
Zoological Journal of the Linnean Society. 201(4), DOI:
10.1093/zoolinnean/zlae090.
"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 Neuquén 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. Zeitschrift für 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
für Geologie und Paläontologie 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 Osmólska (eds.). The Dinosauria Second Edition. University of
California Press. 71-110.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle
Jurassic Cañadón 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
Middle Toarcian, Early Jurassic
Cañadón 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. Moreecently,
Rauhut et al. (2024) used
the Mesozoic Tetrapod Group Theropod Matrix to recover it as either a
basal megalosauroid, basal allosaurian or an allosaurid.
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 Cañadón Asfalto Formation of Argentina highlights phylogenetic
uncertainty in tetanuran theropod dinosaurs. Scientific Reports.
9:18826.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod
dinosaur from the Callovian Balabansai Formation of Kyrgyzstan.
Zoological Journal of the Linnean Society. 201(4), DOI:
10.1093/zoolinnean/zlae090.
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
für Geologie und Paläontologie Abhandlungen. 210(1), 41-83.
Holtz, Molnar and Currie, 2004. In Weishampel, Dodson and Osmólska
(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 Cañadón 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 Crétacé
du nord-est du Bassin de Paris. Bulletin scientifique, historique et littéraire
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 Société
Géologique de France. 4, 439-442.
Sauvage, 1882a. Sur les Reptiles trouvés dans le gault de l'est de la
France. Comptes Rendus
Hebdomadaires des Seances de l'Académie des Sciences. 94, 1265-1266.
Sauvage, 1882b. Recherches sur les reptiles trouvés dans le Gault de
l'est du bassin de Paris. Mémoires de la Société Géologique
de France, série 3. 2(4), 1-42.
Simionescu, 1913. Megalosaurus aus der Unterkreide der Dobrogea. Centralblatt
für Mineralogie, Geologie und Paläontologie. 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 Wüsten
Ägyptens. 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 découverte
et révision systématique: Implications phylogénétiques
et paléobiogéographiques. 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
Piatnitzkysauridae, 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. Rauhut et al. (2024) note that "The
most similar morphology [to Alpkarakush]
is found in Xuanhanosaurus
(IVPP V 6729), which also has well-developed medial and lateral
tubercles on the proximal end of the ventral side of phalanx II-1, in
which, however, the ventral sulcus is less enclosed" and suggested "The
marked similarities between manual phalanx II-1 in Xuanhanosaurus and Alpkarakush
might lend support to a metriacanthosaurid identification of the
former." They recovered it using the Mesozoic Tetrapod Group
Theropod Matrix as a piatnitzkysaurid or metriacanthosaurid.
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 Osmólska (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.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod
dinosaur from the Callovian Balabansai Formation of Kyrgyzstan.
Zoological Journal of the Linnean Society. 201(4), DOI:
10.1093/zoolinnean/zlae090.
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 (DeCourten, 1990)
Aptian, Early Cretaceous
Long Walk Quarry UMNH 0002, Lower Ruby Ranch Member of Cedar Mountain
Formation, Emery County, Utah, US
Material- (UMNH VP 904; = 'UUVP 904') tooth (DeCourten, 1990)
(UMNH VP 4805) tooth (~52x~21x? mm) (DeCourten, 1990)
(UMNH VP 26018) tooth (~68x~33x? mm) (Oswald and Curtice, 2023)
(UMNH VP 26019) anterior tooth (~38x~16x? mm) (Kirkland, Suarez, Suarez
and Hunt-Foster, 2016)
(UMNH VP coll.) several partial teeth, ilium(?) (DeCourten, 1990)
Comments- Discovered between the Summers of 1987 and 1989,
DeCourten (1990) originally reported "Two nearly complete large teeth
(along with some smaller fragments) that belong to an entirely
different dinosaur have been discovered. As much as 12 cm (4') in size
with jagged serrated edges, these teeth are those of a large
carnivorous dinosaur." He stated "it is not possible to identify
the owner of the teeth with precision, but a good candidate seems to be
a dinosaur like Acrocanthosaurus,"
... which lived "in Texas about the same time the Long Walk Quarry
sediments were deposited in Utah, was about the same size, and almost
certainly had teeth of the same general form. For now, we can only
refer to this dinosaur as
Acrocanthosaurus(?)."
DeCourten furthermore said "An ilium (hip blade) has also been
excavated that definitely belongs to a carnivorous dinosaur ... , but
it is far too small to belong to the
Acrocanthosaurus(?)." Finally, DeCourten figures "A large
tooth from a carnivorous dinosaur, probably similar to Acrocantosaurus [sic], from the
Long Walk Quarry. This tooth is over 4 inches long..."
Two very similar publications followed (DeCourten, 1991a, b), with the
shorter one somewhat arbitrarily designated 1991a here since no order
of publication is obvious. DeCourten (1991a) says "several teeth
of a large carnosaur have been recovered from the Long Walk Quarry. The
dagger-like serrated teeth (fig. 5) are about 4 in. (12 cm) long and
slightly elliptical in cross-section. These appear to be the anterior
teeth of a large predator similar to Acrocanthosaurus",
with the same taxonomic conclusions as in 1990. A photo of the
same tooth is now labeled "Large carnosaur tooth from the Long Walk
Quarry. The tooth is more than 4 in. (10 cm) long and probably
represents a theropod similar to Acrocanthosaurus".
DeCourten (1991b) says "two nearly complete teeth, several partial
teeth, and an ilium presently undergoing preparation document the
presence at the Long Walk Quarry of at least one large theropod
dinosaur. The teeth are of the typical dagger-like theropod form, with
serrated edges and a curved anterior margin (Figure 7). The two
complete teeth are 3.3 inches (84mm) and 3.9 inches
(99mm) long, comparable in size to the average tooth of Allosaurus..."
Differently from the 1990 publication however is his statement "The
ilium is at least 16 inches (40cm) long and appears to belong to a
bipedal predator of about average Allosaurus
size as well", so it is no longer considered too small to belong to the
tooth taxon at this point. The same tooth photo is now labeled
"Large theropod tooth (UMNH VP4805) from the Long Walk Quarry" but has
a scale bar for the first time. This indicates a length of ~76
mm, so is apparently the shorter tooth mentioned, but this is also the
total length of which ~32% is root. Based on the figure the crown
height would be ~52 mm instead, but the crown height of the other tooth
is now of course uncertain.
Kirkland et al. (1997) say "Kirkland and Parrish (1995) suggested that
the teeth of the Long Walk Quarry theropod are distinct from Acrocanthosaurus in that they are
much more coarsely serrated", but as the latter meeting abstract only
says that "cf. Acrocanthosaurus"
is part of the middle (Ruby Ranch) fauna perhaps this detail was only
in the associated poster. Kirkland et al. themselves state "an
unidentified large theropod, and Acrocanthosaurus"
are present, listing under "Family Allosauridae ?" "new large theropod"
and "cf. Acrocanthosaurus
sp." in their Table 2, implying some Acrocanthosaurus
was present after all. As Harris (1998) explained "this is in
reference to an isolated tooth (CEU 5107) with very fine serrations
from the Cedar Mountain Formation near the Cleveland-Lloyd Dinosaur
Quarry (J. Kirkland, personal comm., 1998)", which is from the
Price River 3 locality
higher in the formation.
Harris (1998) further says "DeCourten (1991: fig. 5) illustrates a
tooth (UUVP 904) from the Long Walk Quarry in the Cedar Mountain
Formation (Ruby Ranch Member per Kirkland et al., 1997) of Utah
attributed to Acrocanthosaurus
or a similar taxon", referring to what I call DeCourten
1991a.
Levitt-Bussian (pers. comm. 7-2024) clarified UMNH
VP 904 is a Long Walk Quarry theropod tooth, so apparently Harris
accidentally used the number with their older UUVP catalog
system. This indicates it is probably DeCourten's second complete
tooth and Harris was wrong to cite the latter's figure for it. He
continues
"Examination of this specimen indicates that it differs from the Acrocanthosaurus
tooth described by Harris (1997, 1998) in possessing approximately 1
denticle per mm, as opposed to 2 per mm in the Trinity taxon's tooth.
However, precise measurements could not be taken, and examination of
the specimen was impaired by incomplete preparation between the
denticles." DeCourten's (1991b) scale bar suggests 6.3 serrations
per 5 mm at midheight distally, a more commonly used unit of serration
density today. This is indeed much less than the 11.5-17.5 per 5
mm in verified Acrocanthosaurus
teeth. Finally, Harris notes "The association of the ilium in the
same quarry as the teeth may indicate that they originated from the
same taxon, if not the same individual. As no ilium of Acrocanthosaurus
has yet been described, it is impossible to determine if the specimen
represents the Trinity taxon, the taxon represented in the quarry by
the coarsely-serrated teeth, or a different large theropod." Over
twenty-five years later, the ilium of Acrocanthosaurus
remains unknown, but the Long Walk material is all provisionally
assigned to the same unnamed taxon here since Ruby Ranch Acrocanthosaurus
is from a higher level and different locality, all three teeth with
published serration densities have similar values, and DeCourten 1991b
indicated the teeth and ilium were compatible in size. However,
Irmis (pers. comm. 7-2024) reported "I'm pretty sure the ilium turned
out not to be a theropod", so its true identity is yet again uncertain.
Kirkland et al. (2016) note that of the Long Walk elements, "a large
allosaurid [is] noted among the material (figure 29)", with Figure 29B
being labeled "Allosauroid tooth from Long Walk Quarry (UMNH 0002)" and
listed as "large allosauroid". They had earlier cited "the Long
Walk Quarry (UMNH 0002)", showing that to be the locality number not
the specimen number. The figured tooth seems to be mesial, from
either the right dentary or left premaxilla based on the labially
shifted distal carina. At ~90 mm long including the root it
doesn't match either measurement reported by DeCourten (1991b), but is
the same ratio shorter than the longer measurement (90%) as UMNH VP4805
was from the shorter measurement. The much higher specimen number
suggests it is not one of the originally reported teeth, however.
Oswald and Curtice's (2023) SVP poster incorporated two 'Long Walk
Allosaur' teeth- the one figured by Kirkland et al. now identified as
UMNH VP 26019, and other without a root labeled as UMNH VP 26018.
They provide serration measurements of these teeth as 7 mesial per 5 mm
and 7-9 distal per 5 mm.
References- DeCourten, 1990. The Long Walk Quarry: A new horizon
in dinosaur research. Canyon Legacy. 6, 15-22.
DeCourten, 1991a. 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 Geological Society. 19-25.
DeCourten, 1991b. New data on Early Cretaceous dinosaurs from the Long
Walk Quarry and tracksite, Emery County, Utah. Utah Geological
Association Publication 19, 311-324.
Kirkland and Parrish, 1995. Theropod teeth from the Lower and Middle
Cretaceous of Utah. Journal of Vertebrate Paleontology. 15(3), 39A.
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.
Harris, 1998. Large, Early Cretaceous theropods in North America. In
Lucas, Kirkland and Estep (eds.). New Mexico Museum of Natural History
and Science Bulletin. 14, 225-228.
Kirkland, Suarez, Suarez and Hunt-Foster, 2016. The Lower Cretaceous in
east-central Utah - The Cedar Mountain Formation.and its bounding
strata. Geology of the Intermountain West. 3, 101-228.
Oswald and Curtice, 2023. The dragons of Cedar Mountain: Shed teeth
indicate the presence of one or more large allosauroids from the Yellow
Cat member of the Cedar Mountain Formation. 330-331.
undescribed allosauroid (Judd, Irmis and Kirkland, 2013)
Early Albian, Early Cretaceous
near Capitol Reef National Park, Upper Ruby Ranch Member of Cedar
Mountain Formation, Wayne County, Utah, US
Material- (Utah Wn coll.) cervical vertebra, partial sacrum,
partial pelvis, femur
Comments-
Note while Judd et al. (2013) place this in "the lowermost Ruby Ranch
Member", the Albian age makes it part of Kirkland et al.'s (2016) Upper
Ruby Ranch Member.
Judd et al. (2013) stated this "can be placed within Tetanurae based on
a convex anterior face of the presacral vertebrae and a femoral head
oriented dorsomedially", and additionally found this to be an
allosauroid based on- "anterior pleurocoel of the cervical vertebra is
anteroposteriorly elongate, the parapophysis is located in the middle
of the centrum, and there is no articular groove on the proximal
surface of the head of the femur." However, it is similar to
megalosaurids in that "the oblique ligament groove does not extend past
the posterior surface of the femoral head." Kirkland et al. list
this as "Large allosauroid with short neural spines."
References- 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.
Kirkland, Suarez, Suarez and Hunt-Foster, 2016. The Lower Cretaceous in
east-central Utah - The Cedar Mountain Formation.and its bounding
strata. Geology of the Intermountain West. 3, 101-228.
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-Omeñaca, 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 für Geologie und Paläontologie, Abhandlungen. 239(1), 77-99.
Gascó, Cobos, Royo-Torres, Mampel and Alcalá, 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-Omeñaca, Canudo and
Infante, 2005)
Early Barremian, Early Cretaceous
Camarillas Formation, Spain
Material- (MPZ2005/316-317) teeth
Reference- Ruiz-Omeñaca, 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-Omeñaca, 2005)
Early Barremian, Early Cretaceous
Mirambel Formation, Spain
Material- (LAD4r-1) tooth (~22 mm)
Reference- Infante, Canudo and Ruiz-Omeñaca, 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) (Suñer 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 terópodos
del Cretácico Inferior (Fm. Arcillas de Morella, Aptiense) en los yacimientos
del Mas de la Parreta (Morella, Castellón). In Fernández-Martínez
(ed.). XXII Jornadas de Paleontología de la Sociedad Española
de Paleontología. Libro de resúmenes, 124-125.
Suñer and Santos-Cubedo, 2008. Dos dientes de terópodo del yacimiento
ANA, Formación Arcillas de Morella (Aptiense, Cretácico Inferior,
Cinctorres, Castellón). 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 crânien
de dinosaure théropode du Jurassique des Vaches Noires (Normandie, France):
remarques sur la diversité des théropodes jurassiques européens.
Comptes Rendus de l'Academie des Sciences Paris, Série 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 Société géologique 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.
Néraudeau, Allain, Ballèvre, Batten, Buffetaut, Colin, Dabard,
Daviero-Gomez, Albani, Gomez, Grosheny, Le Loeuff, Leprince, Martín-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
Tiourarén 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 Tiourarén
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-
Paul (2016)
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
Osmólska (eds.). The Dinosauria. University of California Press.
169-209.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and
Osmólska (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, Araújo 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 Paléontologie. 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 Osmólska (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 Osmólska (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
Bajocian, Middle Jurassic
Chuanjie 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-
While Wu et al. stated "According to paleostratigraphical studies of
the area, the new dinosaur site is located within the lower member of
the Upper Lufeng Formation (Zhang and Li, 1999)", Sekiya (2011) noted
Fang et al. 2000 restudied the Upper Lufeng Formation and renamed this
lower member the Chuanjie Formation, which has been followed since.
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.
Sekiya, 2011. Re-examination of Chuanjiesaurus
anaensis
(Dinosauria: Sauropoda) from the Middle Jurassic Chuanjie Formation,
Lufeng County, Yunnan Province, southwest China. Memoir of the Fukui
Prefectural Dinosaur Museum. 10, 1-54.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae
(Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2),
211-300.
Alpkarakush Rauhut, Bakirov,
Wings, Fernandes and Hübner, 2024
A. kyrgyzicus
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024
Etymology- "Named after Alpkarakush,
a mythological large bird that often comes to the help of heroes in
critical moments in the 'Manas' epos, one of the central mythological
elements in Kyrgyz culture." "The species epithet refers to the Kyrgyz
Republic, the provenance of the type specimen."
Bathonian-Callovian, Middle Jurassic
FTU-1, Balabansai Formation, Kyrgyzstan
Holotype- (IGB 2-1) (~7-8 m;
~1.250 tons using Campione et al. 2014; >17 year old subadult) left
postorbital (~200 mm tall)
....(IGB 2-2) right postorbital
....(IGB 2-9) left quadratojugal (128 mm tall)
....(IGB 2-10) partial ?twelfth dorsal vertebra (115 mm)
....(IGB 2-11) partial ?thirteenth dorsal vertebra (120 mm)
....(IGB 2-12) partial posterior dorsal neural spine
....(IGB 2-13) fragmentary posterior dorsal neural spine
....(IGB 2-14) fused partial first-fourth sacral vertebrae (430 mm;
s1-s4 ~105-110 mm)
....(IGB 2-15) partial fifth sacral vertebra (110 mm)
....(IGB 2-16) partial dorsal rib
....(IGB 2-17) partial dorsal rib
....(IGB 2-18) partial dorsal rib
....(IGB 2-19) partial dorsal rib
....(IGB 2-20) partial dorsal rib
....(IGB 2-21) numerous dorsal rib fragments
....(IGB 2-22) ?twelfth dorsal neural arch fragment
....(IGB 2-24) left manual phalanx II-1 (80 mm)
....(IGB 2-25) incomplete left ilium
....(IGB 2-26) incomplete right pubis
....(IGB 2-28) partial left pubis
....(IGB 2-29) fused distal pubes
....(IGB 2-30) fused left and distal right ischia (680 mm)
....(IGB 2-31) ilial peduncle of right ischium
....(IGB 2-32) left femur (880 mm)
....(IGB 2-33) right femur (900 mm)
....(IGB 2-34) left tibia (720 mm)
....(IGB 2-35) right tibia (715 mm)
....(IGB 2-36, 2-37) incomplete left fibula
....(IGB 2-38) left astragalocalcaneum (190 mm trans; ast 147, calc 43
mm)
....(IGB 2-39) right astragalocalcaneum
....(IGB 2-40) partial left distal tarsal IV
....(IGB 2-41) right metatarsal II (347 mm)
....(IGB 2-42) right metatarsal III (400 mm)
....(IGB 2-43) left metatarsal III (398 mm)
....(IGB 2-44) right pedal phalanx II-2 (95 mm)
....(IGB 2-45) incomplete right pedal ungual II (~120 mm)
....(IGB 2-46) incomplete pedal ungaul IV (~95 mm)
....(IGB 2-47) incomplete right manual ungual III (~83 mm)
Paratype- (IGB 2-48) (>3
year old juvenile or subadult) right tibia (605 mm)
....(IGB 2-49, 2-50, 2-51, 2-52) incomplete pubes (740 mm)
....(IGB 2-53) partial right ischium
Referred- ...*(IGB 2-3) tooth
(~40x21.5x11 mm)
...*(IGB 2-4) partial tooth
...*(IGB 2-5) partial tooth
?..*(IGB 2-6) anterior tooth (~40x16x14.5 mm)
?..*(IGB 2-7) anterior tooth
?..*(IGB 2-8) anterior tooth
...*(IGB 2-23) incomplete furcula
?..*(IGB 2-27) anterior tooth
Diagnosis- (after Rauhut et
al., 2024; autapomorphies only) posterior dorsal vertebrae with channel
leading from the centroprezygodiapophyseal fossa posteromedially into
pneumatic chambers in the neural arch; manual phalanx II-1 with ventral
sulcus proximally that is almost completely enclosed by medial and
lateral ventral flanges; dorsal margin of ilium slopes steeply
posteroventrally (also in Yangchuanosaurus);
narrow and deep intercondylar groove on anterior side of distal femur;
robust and well-developed medial epicondylar crest on distal femur,
considerably offset proximally from distal end
Comments- Material of the
holotype and paratype was initially discovered in 2006, then excavated
further in 2014, 2016-2017 and 2023. *Rauhut et al. (2024) find
"A few elements, including teeth and a furcula, cannot be confidently
assigned to either of the two individuals. However, as there are no
indications for any other larger vertebrates in the locality, ... these
materials are here regarded as referred specimens. In the case of the
teeth, this should be seen as tentative, especially for the possible
premaxillary teeth, as it cannot be ruled out that these elements
represent shed teeth of scavengers." Unlike other carnosaurs, the
authors note these "have a rather round cross-section, are notably
recurved, and lack carinae or serrations" and "resemble teeth of
certain crocodyliforms (e.g. pholidosaurids)", but since "most Jurassic
terrestrial crocodiles are considerably smaller than the animal that
these teeth were derived from, which is also the case for the vast
majority of the crocodyliform remains described from the Balabansai
Formation" and as "no crocodyliform remains were found at the FTU-1
locality", they "consider the possibility that these teeth represent
premaxillary teeth of Alpkarakush."
IGM 2-27 is in the materials list as part of the holotype pubes and as
a tooth. The fact it is later mentioned as a tooth and is not in
the caption of the pubis figure (Figure 15) suggests it is actually a
tooth. The dorsals IGB 2-10 and 2-11 are labeled "D12(?)" and
"D13(?)" in Figure 8 and positioned with neural spines IGB 2-12 and
2-13 respectively above them, while neural arch fragment IGM 2-22
"might belong to IGB 2-10, but does not directly fit on the broken base
of the centroparapophyseal lamina." While Figure 7 labels
IGB-2-11 "probably last dorsal vertebra", none of these vertebral
positions are defended in the text. As "the vast majority of
elements recovered from site FTU-1" are assigned to the holotype,
Rauhut et al. "consider it likely" the furcula "comes from the same
animal. However, it cannot be excluded that it might represent the
smaller, paratype individual." The metatarsal II is incorrectly
stated to be the left element in the materials list. For the
holotype, Rauhut et al. concluded "The minimum age at the time of death
can be estimated with 17 years. However, possible resorption of
additional growth marks by the expanding medullary cavity during
growth, the weak zonation in the middle unit of the femoral bone wall,
and the slightly higher counts of growth marks in other areas of the
posterolateral bone wall strongly indicate an older age", while the
paratype's tibia only had three growth lines and "belonged to a large
juvenile or a small subadult."
Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix
to recover Alpkarakush as a
metriacanthosaurine in a polytomy with Metriacanthosaurus, Sinraptor and Siamotyrannus. It is placed
as more derived than Shidaisaurus
here due to Carrano et al.'s characters 265-1 (Ilium, height of lateral
wall of brevis fossa relative to medial wall: shorter anteriorly,
exposing medial wall in lateral view), 293-1 (Ischium, shaft
orientation: ventrally curved), 296-1 (Ischium, notch ventral to
obturator process: present) and Rauhut et al.'s mentioned "transverse
dimensions of mid-sacral vertebrae reduced in relation to first and
last sacral", but is more like Shidaisaurus
in Carrano et al.'s character 280-0 (Ilium, shape of posterior margin
of postacetabular process: convex) and intermediate between Shidaisaurus
and derived metriacanthosaurines in Carrano et al.'s character 285
(Pubis, angle between long axes of shaft and boot: 75-90° (0), < 60°
(1)).
Reference- Rauhut, Bakirov,
Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the
Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the
Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.
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. catalog 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 découverte
et révision systématique: Implications phylogénétiques
et paléobiogéographiques. 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 Cañadón 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 Valmitão, 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
Valmitão 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 Andrés,
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 Jurásico Superior
de Valmitão (Lourinhã, Portugal). XXIV Jornadas de la Sociedad
Española de Paleontología, abstracts. 148-149.
Malafaia, Ortega, Escaso, Silva, Ramalheiro, Dantas, Moniz and Barriga, 2009.
A preliminary account of a new Allosaurus individual from the Lourinhã
Group (Upper Jurassic of Torres Vedras, Portugal). Actas de las IV Jornadas
Internacionales sobre Paleontología 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 Neuquén, 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, Río 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 für Geologie
und Paläontologie 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. catalog 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 für Geologie und Paläontologie
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 catalogd 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) concluded "Altispanax dunkeri
(Dames) is a valid form only when the type material is understood to
include only the three dorsal vertebrae described by Owen" (NHMUK
R1828), 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 für Geologie und
Paläontologie - 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 (Lydekker, 1890)
(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. Lydekker (1890) wrote "The
Hollington metatarsus [the incorrectly associated NHMUK R604d and NHMUK
R1525] agreeing, therefore, in relative size with the type tooth [of dunkeri]
and coming from approximatel the same geological horizon, there
is every probability that it belongs to the same species, to which I
accordingly propose to refer it", but Stovall and Langston (1950)
correctly disagreed, stating "It is widely recognized
that the relative size of individual and homologous skeletal elements
cannot be employed as a criterion for generic and specific
identification in the large saurischian reptiles. The conclusion of
Lydekker becomes even more untenable when it is realized that the size
of carnosaurian teeth may vary greatly within the dental series of a
single individual. In view of the above statement, the correlative
geologic horizons seem to be of no great importance in the present
instance inasmuch as the two locations are widely separated."
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. catalog 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.
Lydekker, 1890. Contributions to our knowledge of the dinosaurs of the
Wealden and the sauropterygians of the Purbeck and Oxford Clay.
Quarterly Journal of the Geological Society of London. 46, 36-53.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous
Formations, principally in Europe. Revista del Museo de La Plata. 29,
1-167.
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.
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, Neuquén,
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), Neuquén 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
Etymology- "the name of Acrocanthosaurus atokensis [is] in
allusion to the elongate neural spines."
= "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..
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.
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
Etymology- While not stated by
the authors, atokensis is in
reference to Atoka County where the types were discovered.
= "Acracanthus atokaensis" Langston, 1947
Late Aptian-Early Albian, Early Cretaceous
Antlers Formation, Atoka County, Oklahoma, US
Holotype- (OMNH 10146; = MUO 8-0-S9) (8.5 m, 21+ year old adult)
(skull ~966 mm) left lacrimal, partial left jugal, partial right
postorbital, incomplete right squamosal, fused frontals and parietals
(227 mm), braincase, right ectopterygoid, fragmentary left surangular,
fragmentary left angular, left articular, atlantal intercentrum, axial
fragment, partial third cervical vertebra (96 mm), partial fourth
cervical vertebra (98 mm), incomplete fifth cervical vertebra (123 mm),
incomplete sixth cervical neural arch, partial seventh cervical centrum
(153 mm), partial 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, left coracoid, fused pubes (~838 mm),
left ischium (~621 mm), distal right femur (~949 mm), left tibia (~865
mm), right and fragmentary left fibulae (801 mm), left astragalus
Paratype- (OMNH 10147; = MUO 8-0-S8) (9.5 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), ~seventeenth caudal vertebra,
~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, fused pubes, proximal left femur (~1.052 m),
fragmentary left tibia (~958 mm), left metatarsal II (416 mm), left
metatarsal III (445 mm), left phalanx III-1 (145 mm)
Referred- ?(OMNH 3031) four
gastralia (paleofile.com; OMNH online)
?(OMNH 34878) (juvenile) tooth fragment (OMNH online)
?(OMNH 51788) tooth (25x14.5x7 mm) (Lipka, 1998)
?(OMNH 61249) tooth fragment (OMNH online)
material (Nydam, Cifelli, Brinkman and Gardner, 1997)
Late Aptian-Middle Albian, Early Cretaceous
Antlers Formation, Johnston County, Oklahoma, US
?(OMNH 53487) two teeth, rib fragments, fragments (OMNH online)
Late Aptian-Early Albian, Early Cretaceous
Antlers Formation, Idabel, McCurtain County, Oklahoma, US
(NCSM 14345) (11.5 m, 2.4 or 3.64 tons, adult) skull (1.29 m),
mandibles (1.315 m) (teeth 16.04-93.08x14.42-42.07x8.55x21.44 mm; for
individual measurements see Smith et al., 2005), presacral vertebral
fragments, cervical rib, several partial dorsal ribs, many gastralia
fragments, over fourteen caudal vertebrae (mid caudal ~160, distalmost
preserved 120 mm), six mid chevrons, left scapulocoracoid (scap 970,
cor 210 mm proximodist), right humerus (370 mm), right radius (220 mm),
right ulna (255 mm), right radiale, right intermedium, right semilunate
carpal, right metacarpal I (62 mm), right phalanx I-1 (111 mm),
proximal right manual ungual I, right metacarpal II (116 mm), right
phalanx II-1 (101 mm), proximal right manual ungual II, right
metacarpal III (89 mm), right phalanx III-1 (50 mm), right phalanx
III-2 (42 mm), right femoral shaft (~1.277 m), incomplete right tibia,
partial right astragalus, right calcaneum, right metatarsal I (111 mm),
right phalanx I-1 (70 mm), right pedal ungual I, right metatarsal II
(410 mm), right phalanx II-1 (55 mm), right phalanx II-2 (122 mm),
partial right metatarsal III (~439 mm), right phalanx III-1 (160 mm),
right phalanx III-2 (115 mm), partial right metatarsal IV, right
phalanx IV-1 (85 mm), right phalanx IV-2 (70 mm), right phalanx IV-3
(58 mm), right phalanx IV-4, right pedal ungual IV, right metatarsal V
(200 mm) (Currie and Carpenter, 2000)
....(OMNH 10168) two posterior cervical or anterior dorsal centra,
ischial fragment, distal right femur (Currie and Carpenter, 2000)
Aptian-Late Albian, Early Cretaceous
Cloverly Formation, Big Horn County, Montana, US
?(OMNH 32304) partial tooth (OMNH online)
?(OMNH 60207) tooth (OMNH online)
?(OMNH 60212) tooth fragment (OMNH online)
?(OMNH 60474) tooth (OMNH online)
Aptian-Early Albian, Early Cretaceous
Cloverly Formation, Carbon County, Montana, US
?(OMNH 21958) tooth fragment (OMNH online)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member of the Cloverly Formation,
YPM 64-59, Carbon County, Montana, US
?(YPM 5285) incomplete proximal caudal vertebra (103 mm) (Ostrom,
1970)
Mid-Late Albian, Early Cretaceous
Himes Member of the Cloverly Formation, YPM
63-17, Big Horn County, Wyoming, US
?(UM 20797) partial scapulocoracoid (D'Emic, Melstrom and Eddy, 2012)
Mid-Late Albian, Early Cretaceous
Himes Member of the Cloverly Formation, YPM
63-18, Big Horn County, Wyoming, US
(UM
20796) (502 or 657 kg; 4+ year old juvenile) incomplete dorsal centrum
(~100 mm), incomplete caudal neural
arch (centrum ~75 mm), incomplete pubes, right femur (670 mm), proximal
right fibula, fragments
(Melstrom and D'Emic, 2011; described by D'Emic, Melstrom and Eddy,
2012)
Mid-Late Albian, Early Cretaceous
Himes Member of the Cloverly Formation, YPM
63-19, Big Horn County, Wyoming, US
(YPM 5377) tooth (71x~26x? mm) (Ostrom, 1970)
Mid Albian, Early Cretaceous
Price River 3 / CEU-PR3 / Utah
Em273, Upper Ruby Ranch Member of Cedar Mountain Formation,
Emery County, Utah, US
(CEU 5107) tooth (Kirkland et al., 1997)
Late Albian-Early Cenomanian, Early-Late Cretaceous
Sonorasaurus quarry, Turney
Ranch Formation, Pima County, Arizona, US
(ASDM 330; = ASDM 500-330) tooth (54.7x29x12 mm) (Ratkevich, 1997)
Early Albian, Early Cretaceous
Paluxy Formation of the Trinity Group, Montague County, 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)
Late Aptian-Early Albian, Early Cretaceous
Twin Mountains Formation, Hobson ranch, Texas, US
(SMU 74646) (~9.8 m, 1.87 or 2.80 tons) incomplete jugal,
ectopterygoid,
palatine, partial surangular, articular, partial prearticular, partial
splenial, lateral tooth (95.4x30.7x19 mm), apical 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)
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)
Late Aptian, Early Cretaceous
Dinosaur Park / Cherokee-Sanford Brick Clay Pit / Muirkirk Clay Pit
USNM 41614, Arundel Formation, Prince George's County, Maryland,
US
(USNM 442510; phalanx = USNM 442521 in Martin and Brett-Surman, 1992
unpublished; tooth = USNM 442520 of Carrano, 2024) (associated?) tooth
(~23.6x~16.6x? mm), left pedal phalanx IV-2 (~34 mm) (Martin and
Brett-Surman, 1992 unpublished)
(USNM
466054) (~7 m, 204 kg, 5-6+ year old subadult) partial second or third
dorsal neural arch, fragmentary ~fourth/fifth dorsal neural arch,
incomplete mid caudal vertebra (~75 mm), anterior distal caudal
centrum, distal
caudal centrum (~60 mm), proximal right femur, proximal left tibia
(112x69 mm prox), tibial
fragment, right calcaneum, right phalanx I-1 (45 mm), right proximal
phalanx
II-1, right incomplete pedal ungual II (~64 mm), right phalanx III-1
(78 mm), right phalanx III-2 (57 mm),
?left incomplete pedal ungual III (~73 mm), right phalanx IV-?2
(49 mm), right phalanx IV-?3 (40 mm),
right pedal ungual IV (55 mm) (Martin and Brett-Surman, 1992; described
by
Carrano, 2024)
?(USNM 497718) apical tooth (Lipka, 1998)
?(USNM 497722) tooth (~19.9x~9.2x? mm) (Lipka, 1998)
?(USNM 497723) incomplete tooth (24.5x10x7 mm) (Lipka, 1998)
(USNM 497724) tooth (40x20x10 mm) (Lipka, 1998)
(USNM 497725) right premaxillary tooth (58x~22x13 mm) (Lipka, 1998)
?(USNM 497734) tooth (Frederickson et al., 2018)
?(USNM 497735) partial tooth (Frederickson et al., 2018)
?(USNM 497736) partial tooth (Frederickson et al., 2018)
?(USNM 497741) partial tooth (Frederickson et al., 2018)
(USNM 529426) tooth (~23.9x~15.5x? mm) (Carrano, 2024) 2005
(USNM 540719) incomplete tooth (?x~30.0x? mm) (Carrano, 2024)
Late Aptian, Early Cretaceous
near Miurkirk USNM 41615, Arundel Formation, Prince George's
County, Maryland, US
(USNM 5685; not Goucher College 5685) incomplete tooth (Lull, 1911)
Late Aptian, Early Cretaceous
Arundel Formation, Prince George's County, Maryland, US
?(USNM 535490 in part) (associated?) tooth (~27.3x~11.6x? mm),
partial manual ungual (~72 mm) (Brownstein, 2018)
Diagnoses- (modified from Stovall and Langston, 1950) neural
spines of presacral, sacral and anterior caudal vertebrae more than 2.5
times taller than respective centrum lengths.
(after Currie and Carpenter, 2000) supraoccipital expands on either
side of midline to protrude as double boss behind the nuchal crest;
cervical neural spines have triangular anterior processes that insert
into depressions beneath overhanging processes on preceding neural
spines; accessory process on lateral surface of caudal prezygapophysis.
(after Carrano et al., 2012) absence of nasal extension of antorbital
fossa and associated pneumatopores.
(after D'Emic et al., 2012) femoral head pointed in proximal view.
Other diagnoses- Stovall and
Langston (1950) originally diagnosed Acrocanthosaurus
based on- skull of massive proportions; (supratemporal, preorbital
and/or postorbital?) arcades moderately heavy; orbit and laterotemporal
fenestra somewhat reduced; jugal foramen of antorbital fossa greatly
enlarged; frontals and parietals fused; quadrato-squamosal kinesis
somewhat reduced; cervical centra opisthocoelous; cervical centra of
moderate length; cervical pleurocoels deep and well marginated;
anterior dorsals distinctly opisthocoelous; mid caudals with accessory
neural spines; chevrons closed proximally; chevrons with anterodorsal
processes; pelvis not fused; pubis slender with broadly expanded foot;
ischium straight, slender and elongate; ischium somewhat expanded
distally; tibia strongly bowed laterally; metatarsal III somewhat
constricted proximally.
Currie and Carpenter (2000) also listed the following- lacrimal
contacts postorbital; pleurocoelous fossae and foramina
pronounced on all presacral and sacral vertebrae.
Comments- Langston (1947) originally called this taxon
"Acrocanthus atokaensis" in his thesis before describing it as Acrocanthosaurus
with Stovall in 1950. The initial nomen nudum was published by
Czaplewski et al. (1994). The paratype was discovered in April
1940 on Herman Arnold's farm, while the holotype was found later that
year "three quarters of a mile east on the land of Mr. W. P. Cochran"
(Stovall and Langston, 1950). They first attributed to remains to
the undivided Trinity Sands. The holotype's skull length was
first estimated at 896 mm but NCSM 14345 shows it has a longer snout
than their Figure 2 and scaled to preorbital bar height it would have
been closer to 966 mm. They state "The tenth presacral vertebra
is provisionally identified as the first dorsal", but since "The
parapophyses are set low on the sides of the centrum" it is here
identified as a tenth cervical. Subsequent dorsal number
identifications by those authors have also been reduced by one here to
result in thirteen dorsals. The authors wrote "The estimated
length of the femur in the paratype is 1153 mm", but their table on
page 720 lists this length for the smaller holotype. Using the
complete right femur of SMU 74646 to estimate the holotype's femoral
length from its distal transverse width results in ~949 mm, while the
paratype's distal tibia width being 111% larger would give its femur an
estimated length of ~1.052 m. Note in the 1980s the The Museum of
the University of Oklahoma changed its name to the Oklahoma Museum of
Natural History, and the types were given OMNH numbers to replace their
original MUO numbers. While Currie and Carpenter (2000) claimed "A more
detailed description of the braincase of the holotype has been prepared
by Welles et al." and cited an in prep. paper, with Welles having died
in 1997 and Langston in 2013, this is unlikely to ever be published.
The braincase was CT scanned in 1999 however, and the endocast
described by Franzosa and Rowe (2005). They believed "the fact
that the exoccipital-basioccipital sutures are still unfused suggests
that the specimen was not fully adult." Currie and Carpenter
wrote that unlike the large posterior surangular foramen reported by
Stovall and Langston in the holotype, "it is relatively small (diameter
of 12 mm) in NCSM 14345, where this region is better preserved."
D'emic et al. (2012) found in the holotype the "anterior part of the
fibular thin section reveals 12 LA arranged somewhat unevenly
throughout the outer half of the cortex, and a further nine closely
spaced LAG representing an EFS. Up to 12 LAG may be unaccounted for, as
LAG are not visible in the interior half of the fibular cortex due to
remodeling and the presence of the medullary cavity" and that "The EFS
is interpreted as evidence that this individual was skeletally
mature."
Nydam et al. (1997) recorded Acrocanthosaurus
atokensis
from the McLeod Prison site, later published as Cifelli et al. (1997)
that shows this is OMNH site V706. The remains are likely to be
OMNH 34878 or 61249.
Lipka (1998) lists "a cast of a partial acrocanthosaur lateral tooth,
OMNH 51788, from the Antlers Formation, site V708", which is in Atoka
County according to the OMNH online catalog.
Pittman
(1989) wrote "A recently discovered specimen from Oklahoma
includes the complete skull and jaws, and much of the postcranial
skeleton, including most of the right pes. This specimen is in private
hands and is not currently available for study." Lipka (1998)
followed this up with "A fourth acrocanthosaur skeleton from the
Antlers Formation of Oklahoma is now known. Until recently it had been
in private hands, and little is known about it except that it is nearly
complete and still under study." 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. It was
described in moderate detail by Currie and Carpenter in that
paper. Carrano
(1998) includes measurements of this specimen as "Geol.
Enterprises". Currie and Carpenter used Anderson's femoral
circumference method to estimate its mass as 2.40 tons. The
intermedium was misidentified by Currie and Carpenter as the
ulnare. 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 based on casts made by BHI personnel. They
state "The fourth toe is missing the last two
phalanges" but all phalanges are illustrated in their Figure 14.
D'Emic et al. (2012) determined using histology that "this individual
had just reached skeletal maturity after a lifetime of relatively
continuous growth."
Wyoming specimens- Discovered
in 1963 and initially referred to ?Megalosauridae by Ostrom
(1970), D'Emic et al. (2012) reported regarding YPM 5377 "The size of
this tooth (ca. 8 cm long) resembles those of adult Acrocanthosaurus"
and that it "is also similar to teeth referred to Acrocanthosaurus
atokensis in that it exhibits fine denticulation (11-16
denticles/5 mm;
Harris, 1998b; Smith et al., 2005) and comparable proportions", thus
"This tooth likely pertains to Acrocanthosaurus."
D'Emic
et al. (2012) described UM 20796, collected in 2008, noting "The bones
were found adjacent to the scapulocoracoid of a specimen of Sauroposeidon proteles
(UM 20800...)". Using histology they concluded "Four lines of
arrested growth are visible in the cortex ..., suggesting an age of at
least four years for the specimen. This is a minimum estimate, because
resorption could have resulted in loss of some lines of arrested
growth." D'Emic et al. wrote "Using the equation of Christiansen
and Fariña (2004), the best estimate body mass of UM 20796 at death was 657
kg", while Carrano (2024) estimated a mass of 502 kg "using the
biped formula from Campione et al. (2014)." They also stated
"Other
materials that likely pertain
(but cannot be formally referred) to Acrocanthosaurus
atokensis
from the Cloverly Formation of Wyoming include: YPM 5377, a tooth from
site YPM 63-19 near Lovell, Wyoming; YPM 5285, a caudal vertebra from
site YPM 64-59 near Bridger, Montana; and UM 20797, a partial
scapulocoracoid from near site YPM 63-17 near Lovell."
Utah specimens- The first
supposed Acrocanthosaurus
remains from Utah were two teeth and tooth fragments from the Long Walk
Quarry in the Lower Ruby Ranch Member of the Cedar Mounatin Formation
reported by DeCourten (1990), who said "it is not possible to identify
the owner of the teeth with precision, but a good candidate seems to be
a dinosaur like Acrocanthosaurus,"
... which lived "in Texas about the same time the Long Walk Quarry
sediments were deposited in Utah, was about the same size, and almost
certainly had teeth of the same general form. For now, we can only
refer to this dinosaur as
Acrocanthosaurus(?)." Kirkland et al. (1997) say "Kirkland
and Parrish (1995) suggested that
the teeth of the Long Walk Quarry theropod are distinct from Acrocanthosaurus in that they are
much more coarsely serrated", but as the latter meeting abstract only
says that "cf. Acrocanthosaurus"
is part of the middle (Ruby Ranch) fauna perhaps this detail was only
in the associated poster. However, Kirkland et al. also mention cf.
Acrocanthosaurus sp.
as being present in the Ruby Ranch Member, which Harris (1998)
explains "is in reference to an isolated tooth (CEU 5107) with very
fine serrations from the Cedar Mountain Formation near the
Cleveland-Lloyd Dinosaur Quarry (J. Kirkland, personal comm.,
1998)." Carpenter et al. (2001) mention this find when they say
"The CEU Prehistoric Museum has located a number of other important
sites at the same stratigraphic horizon in the area (Price River
[PR]-2, PR-3, etc.) that have produced ... elements identifiable as
coming from the large theropod Acrocanthosaurus."
Kirkland et al. (2016) more recently mentioned "A large tooth from PR-3
(Utah Em273) [that] has extremely fine serrations for a tooth that size
and seems to be closest to the older genus Acrocanthosaurus."
Note the College of Eastern Utah merged with Utah State University
creating Utah State University Eastern in 2010 and Utah Em273 is a
locality number for the latter, and Price River 3 is close to
Cleveland-Lloyd so this is almost certainly the tooth first mentioned
almost two decades earlier. As Kirkland et al. state "the Price
River Quarries ... have been excavated since the 1990s", CEU 5107 was
discovered between 1990 and 1997.
Arizona tooth- The sauropod
Sonorasaurus began to be
excavated in Spring 1995 in the Turney Ranch
Formation of Arizona, and in 1997 Ratkevich wrote "A single carnosaur
tooth similar to Acrocanthosaurus
(Fig. 7) was found in the quarry and appears to match bite marks found
on the radius and ulna of our skeleton." Figure 7 is labeled
"Carnosaur tooth cf. Acrocanthosaurus
found with the tooth-scared bones of "Sonorasaurus". Length 2.25
inches" showing a tooth in situ. In his official description of
Sonorasaurus, Ratkevich (1998)
mentions "Multiple long bones clearly
exhibit evidence of slash marks from a contemporary carnosaur. The
discovery of an Acrocanthosaurus
tooth associated with the bones allows us to identify at least one of
the flesh-eating species of the Turney Ranch Formation, and may help
account for the relative rarity of ribs, and other skeletal elements,
some of which were likely carried off by this and other
scavengers." However, Scarborough (2000) found "The tooth of a
theropod dinosaur tentatively identified as similar to Acrocanthosaurus
(Figure 5; R. McCord, pers. comm., 1999), was recovered from a block of
quarry discard which has very similar lithology as this massive
sandstone bed ["one meter below the bone bed" of Sonorasaurus].
The strata immediately surrounding the bone bed contain no similar
green claystone clasts, leading to the likelihood that the tooth was
not directly associated with the dinosaur bone bed, but is one meter
beneath, making its association with Sonorasaurus
questionable." His Figure 5 is
lebeled "Tooth recovered from a sandstone bed about one meter below the
bone bed, tentatively identified as close to Acrocanthosaurus, (#500-330, 55 mm
long) with 11 serrations per centimeter." "most
likely position of the Acrocanthosaurus
(?) tooth" is shown in his
Figure 2 of the Sonorasaurus
quarry. McCord and Gillette (2005) noted this as "a theropod tooth,
closely agreeing with and here referred to cf. Acrocanthosaurus".
D'Emic et al. (2016) noted "This tooth was further prepared in 2013"
and are thus able to give its Crown Base Length and Crown Base
Width. Furthermore, they find serration "density can be measured
on the apical part of the distal margin (density=2 denticles/mm), about
the mid-height of the distal margin (2.8 denticles/mm), and mid-height
of the mesial margin (3.4 denticles/mm)", which in the
now standard metric equals 10, 14 and 17 per 5 mm respectively.
This largely falls into the known range of Acrocanthosaurus
(11.5-17.5
distal, 10-20 mesial) and also suggests Scarborough's measurement of 11
serrations was actually per 5 mm, not 1 cm as he wrote. D'Emic et
al. also find "The lateral profile of the tooth when viewed lingually
or labially is weakly sigmoid in outline with a concave basal and
convex apical portion, which was suggested to be diagnostic of
Carcharodontosaurinae (Hendrickx et al., 2015). Based on this
feature, the denticle density, and overall proportions, we refer this
tooth to Carcharodontosaurinae." It's figured as
"Carcharodontosaurinae indet., theropod dinosaur tooth (ASDM 330) from
the mid-Cretaceous Turney Ranch Formation of Arizona,
USA." Scarborough gave the number as 500-330, but as D'Emic et
al. say "All bones of the [Sonorasaurus]
holotype fall under the number ASDM 500", perhaps suggesting 330 was
removed from 500
after Scarborough theorized it derived from a different level of the
quarry than Sonorasaurus.
Oswald and Curtice (2023) include this
tooth in their SVP poster, where they call it ASDM 500:330.
Texas specimens- Langston
(1974) mentioned "a postorbital bone from Wise County, Texas" as Acrocanthosaurus,
and in Carnosauria "a femur a little more than one meter long obtained
from Cretaceous gravels near Bowie, Montague County, Texas. Its size
would suggest Acrocanthosaurus."
He figured a tooth as "Carnosaur, Paluxy Formation. Montague County,
Texas (FMNH-PR965)."
SMU 62271 were said to be similar to Allosaurus, so are
provisionally assigned to Acrocanthosaurus here based on
provenance.
Regarding SMU 74646, Currie and Carpenter (2000) noted that "the
specimen was actually noticed for the first time more than 40 years
earlier" than 1990. It was described in detail by Harris (1998a).
Arundel Acrocanthosaurus? Lipka was
the first author to suggest the presence of Acrocanthosaurus
in the Arundel fauna, with Harris (1998b) noting "Large, isolated
theropod teeth from the Arundel are reportedly (T. Lipka, personal
communication, 1997) virtually identical to those of Acrocanthosaurus..."
Lipka (1998) announced "carnosaur-like teeth [that] now include recent
additions by myself (USNM 497718, 497722, 497723, 497724, 497725 and
497726) and others." These teeth were all found in 1998, and
while briefly described (but not figured) by Lipka who concludes "It
thus seems nearly certain that Acrocanthosaurus
is indicated by the available tooth material", little evidence was
given to support their assignment to the genus. Instead Lipka
vaguely stated "In general, USNM 497725 compares closely with SMU 74646
1-1", "USNM 497724 ... compares closely to OMNH 51788", "USNM 487723
... more closely resembles the Antlers tooth [OMNH 51788]". The
only statement close to involving actual character evidence is "In most
other respects of gross denticle morphology (denticle sizes were not
measured but compared visually) and denticle count (~ 12/cm). USNM
497725 appears to exhibit many of the other features discussed above",
referring to seven dental features here listed under the comments
section "Does Acrocanthosaurus
have diagnostic teeth?" below. Photos from the USNM online
catalog suggest a typo and that Lipka meant to write "~ 12 / 5 mm" in
regard to serration density, which also matches Harris' third
acrocanthosaur dental feature more closely. Brownstein (2018)
figured three "teeth assigned to Acrocanthosaurus
(2-4)" in his Figure 1, which can be identified via the USNM online
catalog as USNM 497722, 497723 and 497724 respectively. He also
figures "a manual ungual possibly assignable to the latter allosaur in
?lateral (5) view" which is part of USNM 535490. The latter was
discovered on March 24, 1999 and also includes a carnosaur-style tooth
and what may be the pubic peduncle of a maniraptoran ilium.
Carrano (2024) stated the presence of "(1) the mesial margin of the
rounded denticles on the mesial carina is subrectangular, with a
flattened surface [character 89], and (2) the mesiodistal axis of the
mid-crown denticles on the distal carina is inclined apically from the
distal margin in lateral view [character 96]" that were found to be
locally diagnostic to the genus by Hendrickx and Mateus (2014)
"indicates that several isolated teeth from the Arundel Clay could
pertain to Acrocanthosaurus,
including USNM 5685, 442520, 497724, 497725, 529426, and 540719; I
assign these to ?Acrocanthosaurus
sp." USNM 5685 is notable as a referred specimen of "Allosaurus" medius
by Lull (1911) that was discovered by Hatcher between October, 1887 and
January, 1888 based on Appendix C in Kranz (1996). Lull says "one
[tooth] ... (No. 5685, Goucher
College) shows decided wear", but Gilmore (1920) correctly notes
this was "Wrongly attributed to Goucher College as this is the catalog
number
of the U. S. National Museum" and this remains true today. Chure
(2000) repeated this error. Lipka (1998) incorrectly states "Lull
(1911b) also reported two other larger crowns under GC 5685" as only
one tooth was reported, matching the catalog. The USNM online
catalog indicates USNM 442520 is actually a turtle shell element from
the Pliocene Yorktown Formation of North Carolina, with Carrano
meaning Arundel theropod tooth USNM 442510 discovered in 1989 (Carrno-
pers. comm. 9-2024). USNM
529426 and 540719 were discovered in 2005 and between 2010-2013
respectively.
Martin and Brett-Surman (1992 unpublished) first reported a USNM
specimen described as "the proximal end of a left tibia, a pedal
phalange [sic] and an ungual" that was said to "still [be] in
preparation
and ha[d] not been cataloged" yet. Identified only as "a
medium-sized
theropod", "the tibia of this specimen resembles those of ornithomimids
and the Late Cretaceous form Dryptosaurus
aquilunguis."
The tibia, pedal phalanx III-2 and pedal ungual II were figured as
"Theropod." Kranz (1996) reported he discovered this in May 1992,
describing the associated remains as "lower leg bones and ribs from
what has been tentatively identified as a large ornithomimid or
"ostrich" dinosaur." Weishampel and Young (1996) wrote "the top
of a
left tibia (fig. 7.7) and a number of toe bones (including one of the
claws) - probably came from an animal 3 to 4 meters (10 to 13 feet)
long" and proposed "Among theropods, the only ones that have flat claws
on the feet and a tibia that has a large crest sticking out the front
are ornithomimosaurs. These are the same features found in the new
Arundel theropod and in Ornithomimus
affinis, so it is reasonable to
conclude that they too are ornithomimosaurs." They concluded the
specimen "perhaps should be called "Ornithomimus"
affinis."
Brownstein
(2018) stated "Gilmore (1920) noted that more material from Arundel
ornithomimosaurs, including a partial tibia, phalanx and pedal ungual
(USNM PAL 466054) had been recovered (Figure 3.1)", referring to this
same specimen and figuring the tibia (incorrectly stated to be in
medial
view- actually lateral) but as Carrano (2024) rightfully stated
"the date of discovery precludes this specimen having been discussed in
Gilmore." Carrano first reported that "Comparative study reveals
that
USNM 466054 is not an ornithomimosaur, but the first Arundel material
that can be confidently identified as Acrocanthosaurus."
He
stated
"Between 1988 and 1997, several other theropod specimens were
discovered in close proximity to the location of USNM 466054. Although
some might pertain to the same individual, they could not be
definitively associated with it are not formally included here" and
exactly which USNM material this refers to has not been
published.
Note the USNM online catalog lists this as being from locality USNM
41615, but Carrano (pers. comm. 7-2024) has confirmed that in the
catalog "for
the moment the two numbers are effectively equivalent." Although
his materials list identifies the figured dorsal as the
second, in the description Carrano is less certain, stating "I
interpret it as pertaining
to dorsal vertebra 2 or 3." Note he identifies two elements as
"right
pedal phalanges III-2 and -3 (or -3 and -4)" but the second could not
be III-4 as that would be an ungual which this second element is
not. The description identifies
these
as III-1 and III-2. Also while his materials list includes "right
pedal unguals II–IV" the description states "Three pedal unguals are
present (Fig. 10), two from the right side and one probably from the
left" with Figure 10 indicating III is the probable left element.
Histologically, Carrano found the "data indicate a minimum age of 5-6
years based on LAGs (though certainly several are missing), and an
animal still undergoing significant growth based on the lack of an
EFS", while size-wise "The available cross-sectional dimensions of the
femur can be used to calculate body mass according to the formula of
Campione et al. (2014), producing an estimated body mass of 204
kg." I estimated its length here as about seven meters, scaling
the six long bone lengths in Carrano's Table 1 to NCSM 14345 and
averaging them to result in 60.3% of the latter's size. This may
be an overestimate however considering general allometric trends in
theropod ontogeny give younger animals longer feet. Carrano noted
that of Weishampel and Young's supposed
ornithomimosaur
characters, that clade does not have notably large cnemial crests,
ventrally flat pedal unguals are typical of non-maniraptoran theropods,
and that furthermore "Distinct from ornithomimosaurs, the pedal unguals
of USNM 466054 lack the ventral “table” and exhibit a low flexor
tubercle rather than medial ventral ridges or fossae." He also
noted
"The anteroposterior narrowness of these [dorsal] neural arches
corresponds well to those of large-bodied theropods such as Allosaurus
(Gilmore, 1920; Madsen, 1976), Acrocanthosaurus
(Harris, 1998),
Mapusaurus (Coria and Currie,
2006) and Tyrannosaurus
(Brochu, 2003)
... unlike the condition in ornithomimosaurs", "an anterior tilt of the
arch relative to the horizontal plane is also visible in the anterior
dorsals of Allosaurus and Acrocanthosaurus (SMU 74646/FWMSH
93B-9;
Harris, 1998), but not the other large-bodied theropods mentioned
above", "Like most allosauroids, the accessory trochanter is distinct
and approximately triangular. The femoral head is elevated and an
oblique articular groove is lacking from its dorsal surface, as in
carcharodontosaurids and megaraptorans ... [It] differs from that of
ornithomimosaurs, which have a horizontally oriented head, and greater
and lesser trochanters that are well separated by a deep incisure whose
depth reaches at least to the ventral margin of the head", and in the
tibia "The lateral proximal condyle is lobular, similar to the
condition in Allosaurus,
carcharodontosaurids, and
megaraptorans."
Finally, "USNM 466054 also possesses one autapomorphy of
Acrocanthosaurus atokensis: a
pointed (i.e., medially narrowing)
femoral head in dorsal view." Carrano "consider[ed] the current
fossil sample of this lineage - in terms of both spatiotemporal density
and specimen completeness - to be insufficient to allow distinguishing
from A. atokensis
(effectively the terminal “node” of the lineage) and
other taxa that might be more closely related to it than to other known
allosauroids (effectively the “stem” of the lineage). Therefore I have
chosen a slightly more ambiguous designation to allow for the
possibility that this specimen might represent another member of the
Acrocanthosaurus lineage" and
so referred to it as "Acrocanthosaurus
cf. A. atokensis".
However I have a more lumping taxonomic
philosophy where given the presence of an autapomorphy and absence of
noted differences, paired with rough stratigraphic equivalence, Arundel
acrocanthosaur specimens are most usefully treated as A. atokensis.
Does Acrocanthosaurus
have diagnostic teeth?
The first teeth associated with an Acrocanthosaurus
specimen are those
in SMU 74646 described by Harris (1998a), who as Lipka (1998) said
"noted features on these two teeth that may prove to be diagnostic for
acrocanthosaurs, which are: 1) small size of denticulation with respect
to the overall size of the tooth, 2) denticle counts of just under 11
denticles per 5 mm midway down each keel, 3) cartouche-shaped distal
denticles with slightly compressed bodies at their bases and denticle
axes perpendicular to the axis of the distal keel, 4) distal denticles
slightly larger (0.6 mm) than mesial denticles (0.5 mm), 5) mesial
denticles are parallelogram-shaped with axes inclined with respect to
the mesial keel becoming more inclined toward the tip, 6)
denticles decrease in size towards the tip, and 7) "apical
denticulation:" contiguous keels with denticles continuing over the
tip." However, the subsequently described complete skull of NCSM
14345 with most dental positions preserved shows a larger variation in
serration density (10-19 per 5 mm mesially, 11.5-17.5 distally) and
DSDI (0.66-1.57). Currie and Carpenter (2000) also stated apical
denticulation "is the same situation in a diverse assemblage of
theropods that includes carcharodontosaurids, velociraptorine
dromaeosaurids, and tyrannosaurids, so this feature has limited
taxonomic utility." Hendrickx and Mateus (2014) recovered two
characters as local apomorphies of Acrocanthosaurus
teeth- "(1) the mesial margin of the rounded denticles on the mesial
carina is subrectangular, with a flattened surface [character 89], and
(2) the mesiodistal axis of the mid-crown denticles on the distal
carina is inclined apically from the distal margin in lateral view
[character 96]" (Carrano, 2024).
Relationships- Stovall and
Langston (1950) stated "the construction of the skull, the general
proportions of the vertebrae, and the broadly expanded pubic "foot,"
the straight elongate shaft of the ischium, and the presence of an
anterior upward projection in the anterior chevrons ... together with
the geographic consideration, seem sufficiently important at present to
justify the incorporation of Acrocanthosaurus
into the family Antrodemidae" instead of Megalosauridae when these were
the only two Jurassic carnosaur (sensu Huene) families recognized
(contra Currie and Carpenter, 2000 who stated it "was assigned to the
Allosauridae" by its describers). Currie and Carpenter 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. Franzosa and Rowe (2005) concluded "Characters
such as the division of the olfactory bulbs and tracts by a mesethmoid,
which occurs in Acrocanthosaurus.
Carcharodontosaurs [sic], and Giganotosaurus, but not in Allosaurus or Sinraptor,
help to strengthen the postulated relationships." When
constrained as an allosaurid in Carrano et al.'s (2012) tetanurine
matrix, it takes 12 additional steps, showing this topology is indeed
improbable.
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Weishampel, 2006. Another look at the dinosaurs of the east coast of
North America. III Jornadas Internacionales sobre Paleontología de
Dinosaurios y su Entorno. 129-168.
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.
Hendrickx and Mateus, 2014.
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.
D'Emic, Foreman and Jud, 2016. Anatomy, systematics, paleoenvironment,
growth, and age of the sauropod dinosaur Sonorasaurus thompsoni from the
Cretaceous of Arizona, USA. Journal of Paleontology. 90(1), 102-132.
Kirkland, Suarez, Suarez and Hunt-Foster, 2016. The Lower Cretaceous in
east-central Utah - The Cedar Mountain Formation.and its bounding
strata. Geology of the Intermountain West. 3, 101-228.
Brownstein, 2018. The biogeography and ecology of the Cretaceous
non-avian dinosaurs of Appalachia. Palaeontologia Electronica. 21.1.5A,
1-56.
Frederickson, Lipka and Cifelli, 2018. Faunal composition and
paleoenvironment of the Arundel Clay (Potomac Formation; Early
Cretaceous), Maryland, USA. Palaeontologia Electronica. 21.2.31A, 1-24.
Oswald and Curtice, 2023. The dragons of Cedar Mountain: Shed teeth
indicate the presence of one or more large allosauroids from the Yellow
Cat member of the Cedar Mountain Formation. 330-331.
Carrano, 2024 (online 2023). First definitive record of Acrocanthosaurus
(Theropoda: Carcharodontosauridae) in the Lower Cretaceous of eastern
North America. Cretaceous Research. 157, 105814.
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, Neuquén, 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 Paleontología 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-Omeñaca, 2004)
Late Cenomanian-Early Turonian, Late Cretaceous
Cerro Lisandro Formation, Río Negro, Argentina
Material- (Endemas PV-2) tooth
teeth
Reference- Canudo, Salgado, Barco, Bolatti and Ruiz-Omeñaca,
2004. Dientes de dinosaurios teropodos y sauropodos de la Formacion
Cerro Lisandro (Cenomaniense superior-Turoniense inferior, Cretacico
superior) en Río 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 Cretácico de Patagonia. Ameghiniana. 36, 17R.
unnamed carcharodontosaurid (Veralli and Calvo, 2003)
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of Rio Neuquén Subgroup, Neuquén, 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, Neuquén, Argentina. Ameghiniana. 40(4), 74R.
Veralli and Calvo, 2004. Dientes de terópodos carcharodontosáuridos
del Turoniano superior-Coniaciano inferior del Neuquén, Patagonia, Argentina.
Ameghiniana. 41(4), 587-590.
undescribed carcharodontosaurid (Martinelli and Forasiepi,
2004)
Campanian-Maastrichtian, Late Cretaceous
Allen Formation, Río Negro, Argentina
Material- (MACN-PV RN 1086) tooth
Reference- Martinelli and Forasiepi, 2004. Late Cretaceous
vertebrates from Bajo de Santa Rosa (Allen Formation), Río 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 São 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 Marília Formation were recognized
as the Serra da Galga Formation.
References- Candeiro, Bergqvist, Novas and Currie, 2004.
Theropod teeth from the Marília 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 Marília
Formation (Late Maastrichtian) at the paleontological site of
Peirópolis in Minas Gerais State, Brazil. Revista Brasileira de
Geociências. 42(2), 323-330.
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
Aptian-Albian, Early Cretaceous
Maortu, Miaogou Formation, 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; absent distal quadrate
groove.
(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.
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
frontals aren't unfused and the axial neural spine is
posterodorsally inclined.
Comments- The material was discovered in 1960. Hao et al. (2024 online) note the neds at Maortu "were mistakenly claimed to belong to "Wulanhushao (Suhongtu) Formation" (Vickaryous et al., 2001), "Dashuigou Formation" (Weishampel et al., 2004) and "Ulansuhai Formation" (Brusatte et al., 2009; Weishampel et al., 2004). However, this fossil-bearing unit ... actually belongs to the Miaogou Formation, which incorporated the formerly abandoned Dashuigou Formation and Wulanhushao Formation (report of Bureau of Geology and Mineral Resources of Nei Mongol Autonomous Region, 1991; Gao et al., 2014)." 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 supposedly related Labocania,
and to
the Tyrannosauroidea based on the highly pneumatized basicranium and
short and deep braincase. Chure found it sister to Labocania
based on "a thick frontal, a lack of a helical groove on the ventral
articular surface of the quadrate, and very slight invasion of the
frontals by the supratemporal fenestra." 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.
More recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group
Theropod Matrix to recover it as a tyrannosauroid outside Megaraptora
plus Eutyrannosauria (note the supplementary information shows a more
precise position was recovered than suggested in Figure 25).
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
Osmólska (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.
Hao, Li, Wang, Wang, Ma, Qinggele, King, Pei, Zhao and Xu, 2024 online. A new oviraptorosaur from the Lower Cretaceous Miaogou Formation of western Inner Mongolia, China. Cretaceous Research. DOI: 10.1016/j.cretres.2024.106023
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod
dinosaur from the Callovian Balabansai Formation of Kyrgyzstan.
Zoological Journal of the Linnean Society. 201(4), DOI:
10.1093/zoolinnean/zlae090.
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). Comunicación 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, Osmólska 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 für 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, São 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 Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîjestufe
(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.
Depéret and Savornin, 1927. La faune de reptiles et de poisons albiens
de Timimoun (Sahara algérien). Bulletin de la société géologique
de France. 27, 257-265.
Stromer, 1931. Ergebnisse der Forschungsreisen Prof. E. Stromers
in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîjestufe
(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 Wüsten
Ägyptens. 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 - aperçu
historique et travaux récents. In Zouhri (ed.). Paléontologie des
vertébrés du Maroc: état des connaissances. Mémoires de la Societé
géologique 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. Géologie et Paléontologie du Gisement de Gadoufaoua
(Aptien du Niger). Cahiers de Paléontologie, 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. Géologie et Paléontologie du Gisement de Gadoufaoua
(Aptien du Niger). Cahiers de Paléontologie, 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. Géologie et Paléontologie du Gisement de Gadoufaoua
(Aptien du Niger). Cahiers de Paléontologie, 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 für Geologie und Paläontologie
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 Faculté 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 séances de la Société
géologique 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, Apesteguía, Gallina,
Mitchell, Smith, Cullen, Shinya, Haluza, Gianechini and Makovicky, 2022
M. gigas
Canale, Apesteguía, Gallina, Mitchell, Smith, Cullen, Shinya, Haluza,
Gianechini and Makovicky, 2022
Late Cenomanian, Late Cretaceous
La Campanas Canyon, Huincul Formation of Rio Limay Subgroup, Neuquén,
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, Apesteguía,
Makovicky, Gallina, Smith, Mitchell, Gianechini and Haluza, 2018. New
Carcharodontosauridae (Dinosauria, Theropoda) from the early Late
Cretaceous of Neuquén, Argentina, yields light on the anatomy of the
group. Reunión de Comunicaciones de la Asociación Paleontológica
Argentina, Libro de Resumenes. R14.
Cullen, Canale, Apesteguía, 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, Apesteguía, 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. Osteología y miología del autopodio posterior en un
nuevo dinosaurio Carcharodontosauridae de la Formación Huincul
(Cenomaniano-Turoniano, Provincia de Neuquén, Argentina). Licenciado en
Paleontología thesis, Universidad Nacional de Río 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 Castaño 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, Cúneo, 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
Cañadón del Gato, Huincul Formation of Rio Limay Subgroup, Neuquén,
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, Río 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). Currie and
Carpenter (2000) mention "new specimens of Giganotosaurus from Argentina" they
used to score the genus in their phylogenetic analysis, which in
hindsight are probably Mapusaurus.
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.
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.
Eberth, Currie, Coria, Garrido and Zonneveld, 2000. Large-theropod
bonebed, Neuquén, 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 Osmólska (eds.). The Dinosauria: Second Edition. 607-613.
Papolio, 2004. Animales Prehistóricos de América 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. Paläontologische 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), Neuquén Province,
Patagonia, Argentina. Cretaceous Research. Journal Pre-proof DOI:
10.1016/j.cretres.2019.104319
Canale, Apesteguía, 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, Neuquén, 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, Río Negro,
Argentina
(MPCA coll.) maxillary tooth (99x42x18 mm) (Valais and Apesteguía, 2001)
Early Cenomanian, Late Cretaceous
? Candeleros Formation of Rio Limay Subgroup, Argentina
(FPDM uncatalogd) tooth (87x44.2x19.5 mm) (Coria and Currie, 2006)
(MMCH coll.) fourteen complete to fragmentary teeth (Neloadino, online
2016)
(MUCP uncatalogd) 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 Shaochilongin
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. Papers such as Currie and Carpenter (2000) provide
additional anatomical details. 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 Apesteguía (2001) briefly described a tooth (MPCA coll.) in an
abstract from the then recently discovered La Buitrera locality of Río 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 uncatalogd
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 Formación Río Limay, Provincia del Neuquén,
Patagonia, Argentina. VII Jornadas Argentinas de Paleontología 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
Paleontología de Vertebrados (1995): Resúmenes de trabajos presentados.
Acta Geological Lilloana. 18(1), 159-160.
Figueiredo, 1998. Os dinossáurios carnívoros: A sua descrição
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, Neuquén - 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.
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.
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 Apesteguía, 2001. Dientes asignables a Giganontosaurus
(Carcharodontososauria, Theropoda) provenientens de "La Buitera",
Formacion Candeleros, provincia de Río 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. Modelización con elementos finitos de un diente referido al género 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 für Geologie und Paläontologie 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 Paleontología 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, Apesteguía, 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.
Gascó, Cobos, Royo-Torres, Mampel and Alcalá, 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 on page 163 stating "The skull of Allosaurus ferox Marsh has an
aperture in the maxillary in front of the antorbital opening. This
aperture is not present in Ceratosaurus",
a maxillary fenestra 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 "The context appears to indicate that "Allosaurus ferox" is a slip of the
pen for Allosaurus fragilis",
which may be true considering Marsh used the nonsensical combination Labrosaurus fragilis in the same
publication (Allosaurus has
priority over Labrosaurus, so
its type species could never be transferred to the latter genus). 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, Osmólska
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 catalog 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 catalog 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 catalog 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, 1896
= 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)
Neotype- (USNM 4734) (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)
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 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 3894) jugal (Currie and Carpenter, 2000)
(UUVP 3943) fifth sacral rib (Kolb, Davis and Gillette, 1996)
(UUVP 3980) femur (630 mm) (Madsen, 1976)
(UUVP 3981) jugal (Currie and Carpenter, 2000)
(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 11) (~6.5 m) material including sacrum, humerus, radius, ulna,
metacarpals and tibia (581 mm) (Currie and Carpenter, 2000; Currie and
Coria, 2016)
(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 uncatalogd/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 uncatalogd) 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 catalog)
premaxilla (Smith et al., 2005)
('YPM 6293'; Orthacanthus specimen in YPM online catalog)
(Chure, 2000a)
(YPM coll.) pubis, incomplete ischium (Marsh, 1879)
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 generally 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. 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 was declared the neotype by the ICZN on
December 29 2023 (ICZN, 2023).
The neotype 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 forelimbs 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.
Marsh (1896) wrote "Pelvis of Labrosaurus
fragilis Marsh, seen from the
left" for Plate XIII Figure 5 which is the pubis and ischium
illustrated by him in 1879 as Allosaurus
fragilis. Nopcsa (1901) listed both the megalosaurid Allosaurus fragilis and the
labrosaurid Labrosaurus fragilis,
apparently believing them to be separate valid species. Hay
(1908) incorrectly stated "Labrosaurus
ferox seems to be called Labrosaurus
fragilis on page 270" because the pelvic elements had never been
referred to ferox before, and
instead Chure (2000) is probably correct in saying "It would appear
that Labrosaurus fragilis is
a lapsus calami for Allosaurus
fragilis." While typos are not generally cataloged here, Labrosaurus fragilis is given an
entry since Nopcsa viewed it as a valid species. Similarly, since
Marsh and Nopsca both listed Allosaurus
fragilis separately, it's not considered a recombination of that
species (which would be impossible anyway as Allosaurus has priority over Labrosaurus).
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 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 the neotype "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 the neotype 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, though it it incomparable to Saurophaganax.
"Madsenius"- This name was orginally reported in a childrens
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 recatalogd 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 neotype 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. Currie and Carpenter (2000) mention "specimens of Allosaurus (MOR 693, TATE 11, USNM
4734) that are 20% smaller than Acrocanthosaurus
have radii, ulnae and metacarpals that are absolutely longer than the
same elements in Acrocanthosaurus",
which makes it possible TATE 11 is the "Wyomingraptor" material, as
allosaur forelimbs are uncommon. Sacral and tibial measurements
were used by Currie and Coria (2016), who also estimated its total
length.
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 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 neotype
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 neotype 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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