Tyrannosauroidea Osborn, 1906 sensu Walker, 1964
Definition- (Tyrannosaurus rex <- Passer domesticus)
(Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019; modified from Sereno, 1998)
Other definitions- (Tyrannosaurus rex <- Ornithomimus velox)
(modified from Padian et al., 1999)
(Tyrannosaurus rex <- Allosaurus fragilis, Ornithomimus velox,
Deinonychus antirrhopus) (Holtz, 2004)
(Tyrannosaurus rex <- Ornithomimus edmontonicus, Troodon
formosus, Velociraptor mongoliensis) (Sereno et al., 2009)
= Deinodontia Flower, 1929
= Deinodontoidea Cope, 1866 emmend. Brown, 1914 sensu Tatarinov, 1964
= Tyrannosauria Olshevsky, 1995
= Tyrannosauroidea sensu Padian et al., 1999
Definition- (Tyrannosaurus rex <- Ornithomimus velox)
= Tyrannosauroidea sensu Holtz, 2004
Definition- (Tyrannosaurus rex <- Allosaurus fragilis, Ornithomimus
velox, Deinonychus antirrhopus)
= Tyrannosauridae sensu Holtz, 2004
Definition- (Tyrannosaurus rex <- Eotyrannus lengi)
= Tyrannosauroidea sensu Sereno et al., 2009
Definition- (Tyrannosaurus rex <- Ornithomimus edmontonicus,
Troodon formosus, Velociraptor mongoliensis)
Ex-Tyrannosauroidea- Lapparent (1960) suggested Carcharodontosaurus
was more closely related to Tyrannosaurus than to megalosaurids/allosaurids,
which has been suggested by several other authors since (Paul, 1988; Kurzanov,
1989; Molnar et al., 1990), though it is now recognized as a carnosaur (Sereno
et al., 1996). Kurzanov also felt that Diplotomodon was a relative of
Carcharodontosaurus, though it has not been restudied recently and is
Theropoda indet. on this site. It was however listed as Tyrannosauroidea indet.
by Holtz (2004) without comment.
When he named it, Walker (1964) included several taxa in Tyrannosauroidea that
are not currently assigned to that clade. Ornithosuchus and Teratosaurus
are crurotarsans (Sereno and Arcucci, 1990; Galton, 1985), while Sinosaurus
is supposedly a synonym of "Dilophosaurus" sinensis (Currie
et al., in prep.). Indosuchus was assigned to Tyrannosauridae, which
was followed by most later authors (e.g. Chatterjee, 1978) until Bonaparte et
al. (1990) determined it is an abelisaurid. Walker also referred Spinosauridae
to his Tyrannosauroidea, but Spinosaurus is a megalosauroid basal tetanurine
(Sereno et al., 1994) and Becklespinax (= Altispinax of Walker)
is a carnosaur (Naish, 1999). Acrocanthosaurus was often thought to be
intermediate between Allosaurus and tyrannosaurids (Walker, 1964; Paul,
1988; Kurzanov, 1989; Bakker et al., 1988), though since being included in cladistic
analyses it is recognized as a carnosaur (e.g. Holtz, 1994).
Ornithomimosaur postcrania are often confused with tyrannosauroids. This includes
numerous Bissekty elements generally referred to Alectrosaurus by Nessov
(1995), but reidentified by Carr (2005) and Sues and Averianov (2015) - manual
ungual CCMGE 431/12457, femora including CCMGE 479/12457 and 724/12457, a tibia,
astragali including CCMGE 447/12457 and 448/12457, metatarsals and pedal unguals
CCMGE 609/12457 and 610/12457.
Eudromaeosaur cranial elements are sometimes confused with tyrannosauroids.
Dromaeosaurus itself was first believed to be a tyrannosaurid (as a deinodontid)
by Matthew and Brown (1922). A more controversial taxon is Itemirus,
which Kurzanov (1976) placed sister to Tyrannosauridae, echoing its placement
in Tyrannosauroidea by Holtz (2004) and Miyashita (2011). Yet Sues and Averianov
(2014) redescribe it as a dromaeosaurid. Nessov (1995) listed the Bissekty maxillary
fragment CCMGE 600/12457 as tyrannosaurid, but Sues and Averianov refer it to
Itemirus.
Chatterjee (1985) described Postosuchus and poposaurids as tyrannosaurid
ancestors. They are now recognized as crurotarsans, with tyrannosaurids being
more closely related to allosaurids than to any Triassic taxon.
Paul (1988) placed several taxa closer to tyrannosaurids than to Allosaurus
in his paraphyletic Allosauridae, which would be defined as tyrannosauroids
using the current definition. Besides Acrocanthosaurus and Indosuchus,
he also included Chilantaisaurus (a more basal avetheropod, even if
megaraptorans are placed in Tyrannosauroidea- e.g. Novas et al., 2013) and Shaochilong
(his Chilantaisaurus maortuensis; which is now considered to be a
carcharodontosaurid- Brusatte et al., 2009). Molnar et al. (1990) thought these
might be tyrannosauroids too, as did Chure (2000) and Holtz (2004) for Shaochilong
but not Chilantaisaurus. Paul also thought Labocania was close
to tyrannosaurids, which is followed by some recent references such as Holtz
(2004) as well. It is tentatively placed as a carcharodontosaurid here. Besides
Paul, Kurzanov (1989) and Molnar et al. (1990) tentatively placed Bahariasaurus
close to tyrannosaurid ancestry, while Chure (2000) assigned it to Tyrannosauridae.
While its position within Orionides is currently unresolved, the tyrannosauroid
characters suggested by these authors are invalid (see entry). Paul also suggested
Erectopus was close to tyrannosaurids, but is is more probably a carnosaur
(Allain, 2002). Finally, Paul placed Indosaurus in a similar position,
but this is an abelisaurid like Indosuchus (Bonaparte et al., 1990).
Several taxa have recently been placed in Tyrannosauroidea, but are here assigned
to other coelurosaur groups. Olshevsky (1995) classified Compsognathus
as a 'tyrannosaurian', largely based on the supposedly didactyl manus. While
compsognathids are similar to some possible basal tyrannosauroids like Dilong,
they are more parsimoniously closer to birds. Buffetaut et al. (1996) originally
assigned Siamotyrannus to Tyrannosauroidea, but is is more probably a
carnosaur (Pharris, DML 1997; Rauhut, 2000). Holtz (2004) assigned Santanaraptor
to Tyrannosauroidea tentatively, and it emerged as a tyrannosauroid more derived
than proceratosaurids and Dilong in Novas et al.'s (2013) and Porfiri
et al.'s (2014) analyses, though it ends up in basal Maniraptora in Dal Sasso
and Maganuco (2011). It is assigned to Compsognathidae here. "Tonouchisaurus" was stated to be a tyrannosauroid
by the press, but has not been described yet and may be another variety of tyrannoraptoran.
Coelurus and Tanycolagreus were both found to be basal tyrannosauroids
by Senter (2007), and the latter was by Porfiri et al. (2014), but may be maniraptoromorphs instead. Bagaraatan was found to be the basalmost
tyrannosauroid by Holtz (2004), and Carr (2005) found it to be sister to Bistahieversor,
but it may be maniraptoromorph as in Rauhut (2000) as well or a chimaera of both clades. Megaraptorans (excluding
Chilantaisaurus and Siats) were found to be coelurosaurs in the
analysis of Novas et al. (2013) and that of Porfiri et al. (2014) which included
the new evidence of Megaraptor's Dilong-like skull.
Tyrannosauroidea defined- Sereno et al.'s (2009) definition is
a revision of Holtz's (2004), substituting Ornithomimus edmontonicus
for O. velox, Velociraptor for Deinonychus, and Troodon
for Allosaurus. Ornithomimus velox is the better ornithomimosaur
specifier, as discussed under Maniraptoriformes. Being the namesake of Deinonychosauria,
Deinonychus is a better specifier than Velociraptor, but Dromaeosaurus
might be better than either due to its priority. Still, I have no problem with
Deinonychus. Replacing Allosaurus with Troodon was a bad
choice, since numerous topologies have had allosaurids sister to tyrannosaurids
but none I'm aware of have had troodontids sister to tyrannosaurids. The only
other taxa that have been suggested to be sister to tyrannosauroids are spinosaurids
(Walker, 1964), carcharodontosaurids (Paul, 1988; Kurzanov, 1989; Molnar et
al., 1990) and compsognathids (Olshevsky, 1995), but the former was explicitly
placed in Tyrannosauroidea and I don't see placing the others in Tyrannosauroidea
as counter-intuitive.
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Brown, 1914. Cretaceous Eocene correlations in New Mexico, Wyoming, Montana.
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Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus
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literature relating chiefly to the year 1928. The Zoological Record, London.
45, 1-77.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du
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Tatarinov, 1964. Nadotryad Dinosauria. Dinozavry. In Orlov (ed.). Osnovy Paleontologii.
12, 523-589.
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.
Kurzanov, 1976. Braincase structure in the carnosaur Itemirus n. gen.,
and some aspects of the cranial anatomy of dinosaurs. Paleontological Journal.
1976, 361-369.
Chatterjee, 1978. Indosuchus and Indosaurus, Cretaceous carnosaurs
from India. Journal of Paleontology. 52(3), 570-580.
Chatterjee, 1985. Postosuchus, a new thecodontian reptile from the Triassic
of Texas and the origin of tyrannosaurs. Philosophical Transactions of the Royal
Society of London, Series B. 309(1139), 395-460.
Galton, 1985. The poposaurid thecodontian Teratosaurus suevicus v. Meyer,
plus reffered specimens mostly based on prosauropod dinosaurs, from the Middle
Stubensandstein (Upper Triassic) of Nordwurttemberg. Stuttgarter Breitage Naturkunde
Ser. B. 116, 1-29.
Bakker, Williams and Currie, 1988. Nanotyrannus, a new genus of pygmy
tyrannosaur, from the latest Cretaceous of Montana. Hunteria. 1, 1-30.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
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Bonaparte, Novas and Coria, 1990. Carnotaurus sastrei Bonaparte, the
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Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
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Sereno and Arcucci, 1990. The monophyly of crurotarsal archosaurs and the origin
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Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
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Sereno, Wilson, Larsson, Dutheil and Sues, 1994. Early Cretaceous dinosaurs
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Nessov, 1995. Dinosaurs of nothern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
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Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku
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Buffetaut, Suteethorn and Tong, 1996. The earliest known tyrannosaur from the
Lower Cretaceous of Thailand. Nature. 381(6584), 689-691.
Sereno, Dutheil, Iarochene, Larsson, Lyon, Magwene, Sidor, Varricchio and Wilson,
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Naish, 1999. Studies on Wealden Group theropods - an investigation into the
historical taxonomy and phylogenetic affinities of new and previously neglected
specimens. Masters thesis. University of Portsmouth. [pp]
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.
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 interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Allain, 2002. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle d�couverte
et r�vision syst�matique: Implications phylog�n�tiques
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Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Sues and Averianov, 2004. Dinosaurs from the Upper Cretaceous (Turonian) of
Dzharakuduk, Kyzylkum Desert, Uzbekistan. Journal of Vertebrate Paleontology.
24(3), 51A-52A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 5(4), 429-463.
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.
Miyashita and Currie, 2009. A new phylogeny of the Tyrannosauroidea (Dinosauria,
Theropoda). Journal of Vertebrate Paleontology. 29(3), 149A.
Sereno, Tan, Brusatte, Kriegstein, Zhao and Cloward, 2009. Tyrannosaurid skeletal
design first evolved at small body size. Science. 326(5951), 418-422.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Dal Sasso and Maganuco, 2011. Scipionyx samniticus (Theropoda: Compsognathidae)
from the Lower Cretaceous of Italy: Osteology, ontogenetic assessment, phylogeny,
soft tissue anatomy, taphonomy, and palaeobiology. Memorie della Societ�
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
281 pp.
Miyashita, 2011. Cranial morphology of the basal tyrannosauroid Itemirus
medullaris and evolution of the braincase pneumaticity in non-avian coelurosaurs.
Journal of Vertebrate Paleontology. Program and Abstracts 2011, 159.
Burch, 2012. Evolution of the forelimb musculature in Tyrannosauroidea (Dinosauria:
Theropoda). Journal of Vertebrate Paleontology. Program and Abstracts 2012,
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Carr, 2012. Ontogeny and phylogeny of cephalic ornamentation in Tyrannosauroidea
(Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology. Program and
Abstracts 2012, 75.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Porfiri, Novas, Clavo, Agnolin, Ezcurra and Cerda, 2014. Juvenile specimen of
Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation.
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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.
Brusatte, Carr, Averianov, Sues, Muir and Butler, 2015. Dinosaur dynasties:
Large theropod turnover in the Mid-Cretaceous as revealed by a new phylogeny
of tyrannosauroids and new fossils from Uzbekistan. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 98.
Sues and Averianov, 2015. Ornithomimidae (Dinosauria: Theropoda) from the Bissekty
Formation (Upper Cretaceous: Turonian) of Uzbekistan. Cretaceous Research. 57,
90-110.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
Currie, Xing, Wu and Dong, in prep. Anatomy and relationships of Sinosaurus
triassicus ("Dilophosaurus sinensis") from the Lufeng Formation
(Lower Jurassic) of Yunnan, China.
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
Aviatyrannis Rauhut, 2003
A. jurassica Rauhut, 2003
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Holotype- (IPFUB Gui Th 1) ilium (~90 mm)
Paratypes- (IPFUB Gui Th 2) fragmentary ilium
(IPFUB Gui Th 3) proximal ischium
?(IPFUB GUI D 89-91) three premaxillary teeth (~6.19 mm) (Zinke, 1998)
Referred- ?(IPFUB GUI D 174-186) thirteen maxillary and dentary teeth
(~10.15 mm) (Zinke, 1998)
Comments- Rauhut (2000) illustrated and briefly described the holotype
ilium as Stokesosaurus sp., before it was described as a new genus by
Rauhut (2003).
Brusatte and Carr (2016) recovered Aviatyrannis as the most basal tyrannosauroid
closer to tyrannosaurids than proceratosaurids in their Bayesian analysis, though
its exclusion from the Dilong+Tyrannosaurus clade is only supported
by 61% posterior probability. Exclusion from Stokesosaurus+Tyrannosaurus
is strong (90%), and from Xiongguanlong+Tyrannosaurus fairly certain
(100%).
References- Zinke, 1998. Small theropod teeth from the Upper Jurassic
coal mine of Guimarota (Portugal). Pal�ontologische Zeitschrift. 72(1/2),
179-189.
Rauhut, 2000. The dinosaur fauna from the Guimarota mine. In Martin and Krebs
(eds.). Guimarota - A Jurassic Ecosystem. Verlag Dr. Friedrich Pfeil. 75-82.
Rauhut, 2003. A tyrannosauroid dinosaur from the Upper Jurassic of Portugal.
Palaeontology. 46, 903-910.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
Proceratosauridae Rauhut, Milner and
Moore-Fay, 2010
Definition- (Proceratosaurus bradleyi <- Allosaurus fragilis,
Tyrannosaurus rex, Coelurus fragilis, Compsognathus longipes, Ornithomimus velox,
Deinonychus antirrhopus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019; modified from Rauhut, Milner and Moore-Fay, 2010)
Other definitions- (Proceratosaurus bradleyi + Kileskus aristotocus)
(modified from Averianov, Krasnolutskii and Ivantsov, 2010)
Comments- Proceratosauridae was originally named by Rauhut et al. (2010)
to include Proceratosaurus and Guanlong, and given a stem-based
definition. Averianov et al. (2010) later found a similar clade (also containing
their new taxon Kileskus) and claimed Rauhut's name was a nomen nudum
as ICZN Article 13.1.1 requires "a description or definition that states
in words characters that are purported to differentiate the taxon." However,
Rauhut et al.'s phylogenetic definition meets that requirement, so Averianov
et al.'s version of Proceratosauridae is unecessary.
References- Averianov, Krasnolutskii and Ivantsov, 2010. A new basal
coelurosaur (Dinosauria: Theropoda) from the Middle Jurassic of Siberia. Proceedings
of the Zoological Institute RAS. 314(1), 42-57.
Rauhut, Milner and Moore-Fay, 2010. Cranial osteology and phylogenetic position
of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from
the Middle Jurassic of England. Zoological Journal of the Linnean Society. 158(1),
155-195.
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
Guanlong Xu, Clark, Forster, Norell,
Erickson, Eberth, Jia and Zhao, 2006
G. wucaii Xu, Clark, Forster, Norell, Erickson, Eberth, Jia and
Zhao, 2006
= Monolophosaurus wucaii (Xu, Clark, Forster, Norell, Erickson, Eberth,
Jia and Zhao, 2006) Paul, 2010
Late Oxfordian, Late Jurassic
TBB 2002, Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Holotype- (IVPP V14531) (~3 m; 12 year old adult) partial skull (345
mm), atlantal intercentrum, third cervical vertebra (65 mm), fourth cervical
vertebra (55 mm), fifth cervical vertebra (51 mm), sixth cervical vertebra (48
mm), seventh cervical vertebra (44 mm), incomplete cervical ribs, first dorsal
vertebra (33 mm), second dorsal vertebra (32 mm), third dorsal vertebra (37
mm), fourth dorsal vertebra (37 mm), fifth dorsal vertebra (35 mm), sixth dorsal
vertebra (37 mm), seventh dorsal vertebra (37 mm), eighth dorsal vertebra (36
mm), ninth dorsal vertebra, tenth dorsal vertebra (47 mm), eleventh dorsal vertebra
(45 mm), twelfth dorsal vertebra (47 mm), thirteenth dorsal vertebra (47 mm),
four dorsal ribs, synsacrum (42, 40, 39, 38, 45 mm), first caudal vertebra (37
mm), second caudal vertebra (36 mm), third caudal vertebra (36 mm), three distal
caudal vertebrae (46, 44 mm), humeri (208, 224 mm), radii (153, 157 mm), ulnae
(175, 178 mm), radiale, intermedium, semilunate carpals, metacarpals I (46, 45 mm),
phalanges I-1 (86, 82 mm), manual unguals I (74, 72 mm straight), metacarpals
II (87 mm), phalanges II-1 (66 mm), phalanges II-2 (78, 74 mm), manual unguals
II (71 mm straight), metacarpals III (one incomplete; 71 mm), phalanges III-1
(32, 29 mm), phalanges III-2 (28 mm), phalanges III-3 (45 mm), manual unguals
III, metacarpal IV (14 mm), ilia (288, 260 mm), pubes (283, 289 mm), ischia
(230 mm), femora (355, 355 mm), tibiae (389, 375 mm), fibulae (360, 364 mm),
astragali, calcanea, distal tarsal, two distal tarsal fragments, metatarsals
I (43, 35 mm), phalanges I-1 (35, 36 mm), pedal unguals I (49 mm), metatarsals
II (185, 188 mm), phalanges II-1 (60, 58 mm), phalanges II-2 (47, 53 mm), pedal
unguals II (one incomplete; 61 mm), metatarsals III (213, 208 mm), phalanges
III-1 (61, 63 mm), phalanx III-2 (50 mm), metatarsals IV (186, 192 mm), phalanges
IV-1 (42, 39 mm), phalanges IV-2 (35, 36 mm), phalanges IV-3 (24, 27 mm), phalanges
IV-4 (20, 22 mm), incomplete pedal ungual IV, partial metatarsals V
Paratype- (IVPP V14532) (~1.2 m; 6 year old juvenile) incomplete skull
(~139 mm), mandibles, cervical vertebrae, cervical ribs, dorsal vertebrae, dorsal
ribs, sacrum, proximal caudal vertebrae, scapula, coracoid, humerus, radius,
ulna, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanges II-1,
phalanx II-2, manual digit III, ilium, partial pubis, femur, tibia, fibula,
metatarsals, pedal phalanges, pedal unguals
Diagnosis- (modified from Xu et al., 2006) distinct opening on the maxilla
close to the premaxilla-maxilla contact; complex, highly pneumatic nasal crest;
low, rugose ridge along the midline of the frontals; dorsally flattened parietal
with two parallel sagittal crests; transverse ridge within the supratemporal
fossa; centropostzygapophyseal lamina on cervicodorsal vertebrae with its dorsal
end expanding laterally; deep, longitudinal sulci on dorsal surfaces of the
distal caudal vertebrae; ventral part of scapular blade with sub-equilateral
triangular cross-section and thick posterior margin; metacarpal II with prominent
medioventral and laterodorsal processes proximally; manual phalanx II-2 with
prominent medioventral process proximally; femoral greater trochanter much narrower
anteroposteriorly than the lesser trochanter; distinct fossa on posterodistal
surfaces of astragalus and calcaneum; pedal phalanx II-1 with prominent paired
ventral processes proximally (also in Aorun).
Other diagnoses- Choiniere (2010) noted the "deep and narrow groove
along the anterior margin of the premaxilla" listed by Xu et al. is also
found in Proceratosaurus and Eotyrannus.
Comments- Discovered in 2002, this is probably the medium-sized "basal tetanuran"
represented by "articulated specimens and a juvenile skull" noted
by Clark et al. (2004). Xu et al. (2006) announced Guanlong and briefly
described the taxon, while the paratype skull and holotype specimen were described
in detail by Choiniere (2010).
Carr (2006) found Guanlong to be sister to Monolophosaurus within
Carnosauria in an unpublished analysis. They were sister taxa based on the shape
of the anterior maxillary process, tall pneumatic, fenestrate sagittal cranial
crest, and obturator foramen in the ischium. While Carr also noted many characters
which differ between the genera, he suggested these may be ontogenetic, as the
taxa are from the same geological unit. However, all other analyses find the
taxa to be separated, with Guanlong either a basal tyrannosauroid or
slightly closer to birds.
References- Clark, Xu, Forster, Wang and Eberth, 2004. New discoveries
from the Middle-to-Upper Jurassic Shishugou Formation, Xinjiang, China. Journal
of Vertebrate Paleontology. 24(3), 78A-79A.
Carr, 2006. Is Guanlong a tyrannosauroid or a subadult Monolophosaurus?
Journal of Vertebrate Paleontology. 26(3), 48A.
Xu, Clark, Forster, Norell, Erickson, Eberth, Jia and Zhao, 2006. A basal tyrannosauroid
dinosaur from the Late Jurassic of China. Nature. 439, 715-718.
Choiniere, 2010. Anatomy and systematics of coelurosaurian theropods from the
Late Jurassic of Xinjiang, China, with comments on forelimb evolution in Theropoda.
PhD thesis, George Washington University. 994 pp.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Stiegler, Choiniere, Xu and Clark, 2012. A multi-element histological analysis
of the Jurassic tyrannosauroid Guanlong wucaii. Journal of Vertebrate
Paleontology. Program and Abstracts 2012, 179.
Sullivan, Yu and Xu, 2014. Manual flexibility and grasping ability in the basal
tyrannosauroid dinosaur Guanlong wucaii. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 237.
Yu, Sullivan and Xu, 2015. Three-dimensional modeling of the manual digits of
the theropod dinosaur Guanlong, with a preliminary functional analysis.
Acta Palaeontologica Sinica. 54(2), 165-173.
Choiniere, Clark, Forster and Xu, in prep.. The anatomy of Guanlong wucaii.
Proceratosaurus Huene, 1926
P. bradleyi (Woodward, 1910) Huene, 1926
= Megalosaurus bradleyi Woodward, 1910
Middle-Late Bathonian, Middle Jurassic
Minchinhampton Reservoir SO 855113, White Limestone, England
Holotype- (NHMUK R4860) partial skull (~290 mm), mandibles (one incomplete;
260 mm), hyoid (110 mm), two cervical rib fragments
Diagnosis- (after Rauhut and Milner, 2008) dorsal process of the premaxilla
inclined slightly anterodorsally and nasal horn core overhanging the premaxillary
internarial bar anteriorly; internarial bar of the premaxilla bifurcating posteriorly
into a posteriorly directed ramus and a dorsally directed ramus; anterior end
of the maxillary antorbital fossa placed considerably anterior and ventral to
the promaxillary fenestra; anteriormost dentary tooth curved anteriorly and
with the carinae oriented labiolingually.
Comments- Woodward (1902) named and described the species in 1910 as
Megalosaurus bradleyi. It was placed in Megalosaurus as it has
four premaxillary teeth, unlike Ceratosaurus - the other 'megalosaurid'
mentioned by Woodward. Huene (1926) thought that this species was most closely
related to Ceratosaurus because of the nasal crest and therefore separated
it from Megalosaurus as Proceratosaurus bradleyi. He distinguished
it from Ceratosaurus by the greater amount of premaxillary and maxillary
teeth, as well as labially fluted premaxillary teeth. He distinguished it from
'megalosaurs' (Megalosaurus, Eustreptospondylus and Allosaurus)
by the shape of the external naris, the shape and breadth of the dorsal maxillary
process, the height of the antorbital fenestra and the lower mandibular joint.
Huene (1926) later formalized the relationship between Proceratosaurus
and Ceratosaurus by placing them both in the Ceratosauridae. Paul (1988)
was the first to suggest Proceratosaurus is a coelurosaur, specifically
related to Ornitholestes in Ornitholestinae within the Allosauridae.
This was based on their procumbant anterior dentary teeth, conical anterior
teeth (both common in basal coelurosaurs), small anterior teeth and nasal horn
(both not present in Ornitholestes). Holtz (2000) recovered Proceratosaurus
as a coelurosaur more basal than Ornitholestes, though he later (Holtz,
2001; Holtz et al., 2004) recovered it as a sister taxon of Ornitholestes.
Rauhut (2003) recovered it as the most basal coelurosaur however. Naish (online,
2006) noted his unpublished thesis finds Proceratosaurus to be a basal
tyrannosauroid. Rauhut and Milner (2008) restudied the skull and referred Proceratosaurus
to Tyrannosauroidea based on- short premaxilla; well-developed jugal recess;
steeply sloping basisphenoid; premaxillary teeth that are considerably smaller
than the maxillary teeth; D-shaped anteriormost premaxillary teeth. This was
expanded on in Rauhut et al. (2010).
References- Woodward, 1910. On a skull of Megalosaurus from the
Great Oolite of Minchinhampton (Gloucestershire). Quarterly Journal of the Geological
Society of London. 66(262), 111-115.
Huene, 1926. On several known and unknown reptiles of the order Saurischia from
England and France. Annal and Magazine of Natural History. ser. 9. 17, 473-489.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista del Museo de La Plata. 29, 35-167.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: The horned theropods
of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus
- and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River
- Deinonychus, Velociraptor and Saurornitholestes. Hunteria.
2(4), 1-9.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Holtz. 2000. A new phylogeny of the carnivorous dinosaurs. GAIA. 15, 5-61.
Rauhut. 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osm�lska
(eds.). The Dinosauria (second edition). University of California Press, Berkeley.
71-110.
Naish, online 2006. http://darrennaish.blogspot.com/2006/07/war-on-parasites-oviraptorosaurs-eye.html
Rauhut and Milner, 2008. Cranial anatomy and systematic position of the Middle
Jurassic theropod dinosaur Proceratosaurus from England. Journal of Vertebrate
Paleontology. 28(3), 130A.
Rauhut, Milner and Moore-Fay, 2010. Cranial osteology and phylogenetic position
of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from
the Middle Jurassic of England. Zoological Journal of the Linnean Society. 158(1),
155-195.
Kileskus Averianov, Krasnolutskii
and Ivantsov, 2010
K. aristotocus Averianov, Krasnolutskii and Ivantsov, 2010
Bathonian, Middle Jurassic
Upper Itat Formation, Russia
Holotype- (ZIN PH 5/117) incomplete maxilla (~290 mm)
Paratypes- ....(ZIN PH 6/117) premaxilla
....(ZIN PH 7/117) posterior surangular
....(ZIN PH 8/117) metacarpal II (110.9 mm)
....(ZIN PH 9/117) manual phalanx II-1 (74.3 mm)
....(ZIN PH 10/117) metatarsal I (46.6 mm)
....(ZIN PH 11/117) metatarsal III (219 mm)
....(ZIN PH 12/117) pedal phalanx II-2 (37.5 mm)
....(ZIN PH 13/117) pedal ungual III
Referred- ? distal caudal vertebrae (Averianov, Krasnolutskii and Ivantsov,
2010)
Diagnosis- (after Averianov et al., 2010) ascending process of maxilla
confluent with anterior rim of maxilla and gently sloping anterodorsally.
Comments- Both Averianov et al. (2010) and Brusatte et al. (2010; and
future varients) find this to be the most basal proceratosaurid.
References- Averianov, Krasnolutskii and Ivantsov, 2010. A new basal
coelurosaur (Dinosauria: Theropoda) from the Middle Jurassic of Siberia. Proceedings
of the Zoological Institute RAS. 314(1), 42-57.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Sinotyrannus Ji, Ji and
Zhang, 2009
S. kazuoensis Ji, Ji and Zhang, 2009
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (KZV-001) (~9-10 m) anterior skull (~1 m), anterior dentaries,
partial dorsal vertebra, dorsal vertebra, incomplete dorsal vertebra, incomplete
dorsal vertebra, incomplete phalanx II-1, phalanx II-2, manual ungual II, ilia
(one incomplete, one partial; 770 mm), proximal pubes, ischial fragments
Comments- Brusatte et al. (2010) found this to be a proceratosaurid,
sister to Proceratosaurus itself.
References- Ji, Ji and Zhang, 2009. First large tyrannosauroid theropod
from the Early Cretaceous Jehol Biota in Northeastern China. Geological Bulletin
of China. 28(10), 1369-1374.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Yutyrannus Xu, Wang, Zhang, Ma,
Xing, Sullivan, Hu, Cheng and Wang, 2012
Y. huali Xu, Wang, Zhang, Ma, Xing, Sullivan, Hu, Cheng and Wang,
2012
Barremian-Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Holotype- (ZCDM V5000) (adult; 1400 kg) incomplete skull (~905 mm), incomplete
mandible, presacral vertebrae including tenth to thirteenth dorsal vertebrae,
dorsal ribs, gastralia, synsacrum, first to thirty-first caudal vertebrae, chevrons,
scapulae (600 mm), coracoids, humerus, radius (273 mm), ulna, metacarpal I,
metacarpal II, metacarpal III (150 mm), manual phalanges, manual unguals, partial
ilia (~710 mm), incomplete femur (850 mm), tibiae (725 mm), fibulae, astragalaus,
distal tarsal III, distal tarsal IV, metatarsals II, phalanx II-2, pedal ungual
II, metatarsal III (350 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal
ungual III, metatarsals IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx
IV-4, pedal ungual IV, pedal phalanges, pedal unguals, metatarsal V, feathers
Paratypes- (ELDM V1001) (subadult; 600 kg) skull (630 mm), mandible,
hyoids, atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth
cervical vertebra, sixth cervical vertebra, partial seventh cervical vertebra,
cervical ribs, twelfth dorsal centrum, thirteenth dorsal centrum, posterior
dorsal ribs, sacrum, first caudal vertebra, incomplete ilium (~530 mm), pubis,
proximal ischium, femur (613 mm), tibia (623 mm), fibula, astragalus, calcaneum,
distal tarsal IV, metatarsal II, metatarsal III (312 mm), incomplete phalanx
III-1, metatarsal IV, phalanx IV-1, feathers
(ZCDM V5001) (subadult; 500 kg) skull (~800 mm), mandibles, atlas, axis, eight
cervical vertebrae, cervical ribs, nine dorsal vertebrae, dorsal ribs, gastralia,
sacrum, twenty-two caudal vertebrae, several chevrons, scapulae (510 mm), coracoids,
humerus, radius (220 mm), ulna, carpals, metacarpals I, phalanges I-1, manual
unguals I, metacarpals II (130 mm), phalanges II-1, phalanges II-2, manual unguals
II, metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3, manual
unguals III, ilia (620 mm), pubes, ischia, femora (650 mm), tibia (655 mm),
fibula, astragalus, calcaneum, phalanx I-1, metatarsals II, phalanges II-1,
phalanx II-2, pedal ungual II, metatarsals III (350 mm), phalanges III-1, phalanx
III-2, phalanx III-3, pedalm ungual III, metatarsals IV, phalanx IV-1, phalanx
IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, feathers
Diagnosis- (after Xu et al., 2012) rugose, highly fenestrated midline
crest formed by the premaxillae and nasals; anteroventrally projecting orbital
process in the area of the junction between the frontal and jugal processes
of the postorbital; large concavity on the lateral surface of the main body
of the postorbital; external mandibular fenestra located mostly within the surangular.
differs from Sinotyrannus in- morphologically lateral surface of the
maxillary process of the premaxilla faces dorsally; maxilla lacks an anterior
ramus; maxillary fenestra is posteriorly positioned; antorbital fossa has a
posteroventrally sloping ventral margin; ilium has a straight dorsal margin
and a postacetabular process whose ventral margin bears a lobe-like flange.
Comments- The type material was acquired from a fossil dealer, making
its exact provenence uncertain. Xu et al. added it to Carr's tyrannosauroid
analysis and found it emerge sister to Eotyrannus, Xiongguanlong
and more derived tyrannosauroids. Brusatte et al. (2015) found it to be a proceratosaurid
in a larger unpublished analysis, published the next year as Brusatte and Carr
(2016).
References- Xu, Wang, Zhang, Ma, Xing, Sullivan, Hu, Cheng and Wang,
2012. A gigantic feathered dinosaur from the Lower Cretaceous of China. Nature.
484, 92-95.
Brusatte, Carr, Averianov, Sues, Muir and Butler, 2015. Dinosaur dynasties:
Large theropod turnover in the Mid-Cretaceous as revealed by a new phylogeny
of tyrannosauroids and new fossils from Uzbekistan. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 98.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
Pantyrannosauria Delcourt and Nelson Grillo, 2018
Definition- (Tyrannosaurus rex, Dilong paradoxus <- Proceratosaurus bradleyi) (Delcourt and Nelson Grillo, 2018)
Reference- Delcourt and Nelson Grillo, 2018. Tyrannosauroids from the Southern
Hemisphere: Implications for biogeography, evolution, and taxonomy.
Palaeogeography, Palaeoclimatology, Palaeoecology. 511, 379-387.
DOI:10.1016/j.palaeo.2018.09.003
Calamosaurus Lydekker, 1891
= Calamospondylus Lydekker, 1889 (preoccupied Fox in Anonymous, 1866)
C. foxi (Lydekker, 1889) Lydekker, 1891
= Calamospondylus foxi Lydekker, 1889
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (NHMUK R901) (~3.5 m; subadult) anterior cervical vertebra (40
mm), anterior cervical centrum
Referred- (IWCMS 2015 coll.) cervical centrum (Isle of Wight Council,
2015)
Diagnosis- (after Naish et al., 2001) anterior cervical transverse processes
square in section.
Comments- Lydekker named this taxon Calamospondylus without realizing
Fox had previously used the name for a different specimen (Calamospondylus
oweni). It cannot be compared with Calamospondylus or Aristosuchus,
so synonymy with either is inappropriate. Naish (online, 2006; Naish and Martill,
2007; 2011) correctly realized Calamosaurus' cervicals are extremely
similar to Dilong's, suggesting it's also a tyrannosauroid. In a 2004
DML post, Naish also states "there are presently three soon-to-appear papers
on Calamosaurus and its referred material." Two may be his 2011
theropod chapter and/or 2007 British saurischian review with Martill, but another
is a short paper detailing the similarity between Calamosaurus and Dilong
(Naish, online 2006) which remains unpublished. A cervical centrum donated to
the IWCMS collections in 2015 seems referrable to Calamosaurus (Isle
of Wight Council, 2015), but has yet to be described.
The tibia (NHMUK R186) often referred to this species cannot come from the holotype
individual due to size disparity and is not comparable to the holotype in any
case (Naish et al., 2001), though it may be an ornithomimosaur (Allain et al.,
2014). Naish (DML, 2002) reported a new specimen referred to Calamosaurus,
which has since been placed on display at the Dinosaur Expeditions Centre (cast
UOP-C002-2004). It consists of a dorsal vertebra, distal tibia and proximal
metatarsal (Mattsson, pers. comm. 2015), which are undescribed. The tibia was
figured by Naish (2011) and said to be "morphologically identical"
to NHMUK R186, so is listed with that specimen here. A second undescribed specimen
referred to Calamosaurus was discovered in 2014 and is also on display
there (Mattsson, pers. comm. 2015). Based on apparent similarity between the
tibiae of the 2002 and 2014 specimens, they are listed in the same entry here.
References- Lydekker, 1889. On a coelurid dinosaur from the Wealden.
Geological Magazine, decade 3. 4(297), 119-121.
Lydekker, 1891. On certain ornithosaurian and dinosaurian remains. Quarterly
Journal of the Geological Society of London. 47, 41-44.
Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: Theropods. In Martill
and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association.
242-309.
Naish, DML 2002. https://web.archive.org/web/20201114071259/http://dml.cmnh.org/2002Apr/msg00529.html
Naish, DML 2004. https://web.archive.org/web/20160806140122/http://dml.cmnh.org/2004Oct/msg00236.html
Naish, online 2006. http://darrennaish.blogspot.com/2006/06/basal-tyrant-dinosaurs-and-my-pet.html
Naish and Martill, 2007. Dinosaurs of Great Britain and the role of the Geological
Society of London in their discovery: Basal Dinosauria and Saurischia. Journal
of the Geological Society, London. 164, 493-510.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria,
Theropoda): An undetected record. Geologica Acta. 12(2), 127-135.
Isle of Wight Council, 2015. Another important dinosaur fossil discovery made
on the Isle of Wight. On the Wight. http://onthewight.com/2015/04/08/another-important-dinosaur-fossil-discovery-made-on-the-isle-of-wight/
Dilong Xu, Norell, Kuang, Wang,
Zhao and Jia, 2004
D. paradoxus Xu, Norell, Kuang, Wang, Zhao and Jia, 2004
Late Valanginian-Hauterivian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V14243) (1.6 m; subadult) incomplete skull (166 mm),
mandibles, skeleton including mid cervical vertebra (24 mm), posterior dorsal
vertebra (30 mm), dorsal ribs, gastralia, twelve posterior caudal vertebrae
(19-29 mm), scapula (69 mm), coracoid (48 mm), humerus (96 mm), radius (~90
mm), ilium (138 mm), pubis (134 mm), femora (181 mm; one partial), tibiae (203
mm), fibula (196 mm), metatarsal II (112 mm), metatarsal III (117 mm), metatarsal
IV (111 mm), pedal phalanges
Paratypes- (IVPP V14242) (1.3 m; subadult) incomplete skull (132 mm),
presacral vertebrae including mid cervical vertebra, pedal phalanx I-1, pedal
ungual I, metatarsal II, phalanx II-1, metatarsal III, phalanx III-1, metatarsal
IV
(TNP01109) partial skull
Early Aptian, Early Cretaceous
Jianshangou Bed of Yixian Formation, Liaoning, China
Referred- (IVPP V11579) (1.5 m; subadult) (skull ~158 mm) maxillae (96
mm), lacrimal, partial jugal, postorbital, squamosal, splenial (69 mm), surangular
(95 mm), incomplete dentary, teeth, atlas (22 mm), axis (21 mm), six cervical
vertebrae, three cervical ribs, first dorsal vertebra (26 mm), second dorsal
vertebra (21 mm), third dorsal vertebra (21 mm), fourth dorsal vertebra (26
mm), fifth dorsal vertebra (26 mm), sixth dorsal vertebra (23 mm), seventh dorsal
vertebra (22 mm), three dorsal vertebrae, five dorsal ribs, gastralia, first
caudal vertebra (26 mm), second caudal vertebra (25 mm), third caudal vertebra
(26 mm), fourth caudal vertebra (26 mm), fifth caudal vertebra (25 mm), sixth
caudal vertebra (26 mm), seventh caudal vertebra (23 mm), eighth caudal vertebra
(26 mm), ninth caudal vertebra (25 mm), ten caudal vertebrae, nine chevrons,
scapula (89 mm), coracoid (~48 mm), three sternal ribs, distal humerus, distal
radius, metacarpal I (21 mm), phalanx I-1, manual ungual I (39 mm), metacarpal
II (43 mm), phalanx II-1 (28 mm), phalanx II-2 (43 mm), manual ungual II, metacarpal
III (33 mm), phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III
(24 mm), partial ilium, pubis, ischium, femur, tibiae, partial fibula, metatarsal
I (28 mm), metatarsal II (109 mm), phalanx II-1, phalanx II-2, pedal ungual
II, metatarsal III (118 mm), phalanx III-1, phalanx III-2, pedal ungual II,
metatarsal IV (108 mm), metatarsal V (42 mm), feathers (Xu, Norell, Kuang, Wang,
Zhao and Jia, 2004)
Diagnosis- (modified from Xu et al., 2004) two large pneumatic recesses
dorsal to the antorbital fossa on the maxilla; extremely long descending process
of the squamosal extending close to the mandibular articulation of the quadrate;
lateral projection of the basisphenoid anterior to the basal tuber; very deep,
sub-circular interspinous ligamentous fossae on cervical vertebrae; robust scapula
with a wide distal end (distal end twice as wide as the proximal scapular blade);
hypertrophied coracoid (dorsoventral length about 70% of the scapular length).
Comments- IVPP V11579 may be a separate species. White (2009) illustrated
and described a Dilong
metatarsus, labeled IVPP V11579 in the text but V14242 in the figure.
The figure is probably correct, as the metatarsus in IVPP V11579 is
articulated differently. The endocast was described by Brusatte
et al. (2016), and published in detail as Kundrat et al. (2020).
While many cladistic analyses have found this to be a basal tyrannosauroid,
some recent examples find it to be slightly closer to birds.
References- Xu, Norell, Kuang, Wang, Zhao and Jia, 2004. Basal tyrannosauroids
from China and evidence for protofeathers in tyrannosauroids. Nature. 431, 680-684.
White, 2009. The subarctometatarsus: Intermediate metatarsus architecture demonstrating
the evolution of the arctometatarsus and advanced agility in theropod dinosaurs.
Alcheringa. 33(1), 1-21.
Janacek, Guo, Xu and Kundrat, 2014. 3D reconstruction of the endoneurocranial
shape of a basal tyrannosauroid. Journal of Vertebrate Paleontology. Program
and Abstracts 2014, 153.
Brusatte, Muir, Averianov, Balanoff, Bever, Carr, Kundrat, Sues,
Williamson and Xu, 2016. Brains before brawn: Neurosensory evolution in
tyrannosauroid dinosaurs. Journal of Vertebrate Paleontology. Program
and Abstracts, 106.
Kundr�t, Xu, Hančov�, Gajdoš, Guo and Chen, 2020 (online 2018).
Evolutionary disparity in the endoneurocranial configuration between
small and gigantic tyrannosauroids. Historical Biology. 32(5), 620-634.
Nuthetes Owen, 1854
N. destructor Owen, 1854
= Megalosaurus destructor (Owen, 1854) Steel, 1970
Middle Berriasian, Early Cretaceous
Cherty Freshwater Member of Lulworth Formation of Purbeck Limestone Group, England
Holotype- (DORCM G 913) (~2.3 m) (skull ~250 mm) partial dentary, teeth
Referred- (NHMUK 48207) dentary fragments, teeth (Owen, 1878)
(NHMUK 48208) several teeth (Owen, 1878)
?(NHMUK R 15870) maxillary tooth (15.5 mm) (Owen, 1879)
(NHMUK R 15871) premaxilary tooth (7 mm) (Owen, 1879)
(NHMUK R 15872) dentary tooth (5 mm) (Owen, 1879)
(NHMUK R 15873) dentary tooth (5 mm) (Owen, 1879)
(NHMUK R 15874) maxillary tooth (8.25 mm) (Owen, 1879)
(NHMUK R 15875) maxillary tooth (8.5 mm) (Owen, 1879)
(NHMUK R 15876) dentary tooth (1.5 mm) (Owen, 1879)
(NHMUK R 15877) lateral tooth (5 mm) (Milner, 2002)
?(NHMUK R 15878) maxillary tooth (16 mm) (Milner, 2002)
Early Berriasian, Early Cretaceous
Marly Freshwater Member of Lulworth Formation of Purbeck Limestone Group, England
(CAMSM J13951) lateral tooth (Milner, 2002)
Berriasian, Early Cretaceous
Purbeck Limestone Group, England
? material (Delair, 1959; Benton and Spencer, 1995)
Description- Initial comparisons are to dromaeosaurids, as Milner (1999,
2002) proposed Nuthetes was a member of this clade. The holotype anterior
dentary fragment suggests a cranial length of ~250 mm, and a total length of
2.3 meters, scaling from Deinonychus.
The dentary has an upper row of foramina, like dromaeosaurids and most other
theropods, which are circular anteriorly (unlike Sinornithosaurus). A
lower row seems to be absent, replaced by a groove that runs posterodorsally
until fading around the seventh tooth. This seems unique within dromaeosaurids.
Separate interdental plates are present, like Sinornithosaurus (in the
anterior dentary at least) and Bambiraptor. The symphysis is represented
by horizontal striations. The Mackelian groove is reduced to a poorly defined
shallow depression terminating at the fifth tooth, with a foramen directly anterior.
In Deinonychus and Saurornitholestes, the groove is shallow but
well defined and the foramen located at the third or fourth tooth. Velociraptor's
looks similar, but may end a bit more posteriorly, while Dromaeosaurus'
ends more anteriorly, at the second tooth or so. Sinornithosaurus' extends
to the third tooth, but is deep and narrow.
Teeth (numbering >9) are typically theropod in several respects; they are
laterally compressed, recurved and serrated. The tip of some (all?) recurved
teeth is located posterior to the alveolar edge, which is seen in dromaeosaurids.
Milner reports the extent of serration on mesial carinae varies from the apical
third to the whole crown, while the distal serrations are restricted to the
apical half when present at all. I'm unaware of this occurring in any other
theropod, as most lose their mesial serrations before their distal ones. And
indeed one can see in plate 1 (6, 10) that distal serrations extend to the base
of the crown in at least some teeth. Thus, I think Milner confused mesial and
distal a few times in the text. What I suspect is really the case is that the
mesial carinae are serrated for up to half their length (sometimes unserrated)
while the distal carinae are completely serrated. This is similar to "velociraptorines"
and Sinornithosaurus, but distinct from Dromaeosaurus (whch seemingly
always has serrated mesial carinae) and Tsaagan, Microraptor and
Cryptovolans (which lack mesial serrations). Heterodonty is observed,
as 40% of the teeth are longer and more slender than the rest. This seems more
developed than in Saurornitholestes. Constriction between the crown and
root is comparable to Saurornitholestes, so is much less than in Microraptor.
The DSDI varies from 1.14 to 1.55, which is almost identical to Sinornithosaurus
(1.13-1.43), larger than dromaeosaurines (.81-1.18) and comparable to the low
end of "velociraptorines" (1.19-2.33). Serrations are smaller than
in most dromaeosaurids (4.5-8/mm compared to 2.5-2.8/mm in Dromaeosaurus,
2.4-2.6/mm in Utahraptor, 4-5/mm in Saurornitholestes, 3.2-3.6/mm
in Deinonychus, 5/mm in Bambiraptor and Velociraptor; 3-3.6/mm
in Achillobator) except for Sinornithosaurus and Microraptor,
which have 7-14/mm and 8/mm respectively. Serrations have blunt rounded tips,
unlike in "velociraptorine" and some anterior Sinornithosaurus
teeth. Dromaeosaurines, Microraptor and most Sinornithosaurus
teeth share this plesiomorphy with Nuthetes. Some serrations are asymmetrical,
being almost hooked apically. Sinornithosaurus has only perpendicular
serrations, while "velociraptorines'" and Microraptor's are
hooked. Dromaeosaurus shows similar variation to Nuthetes.
A premaxillary tooth is said to be serrated only mesially, which is also seen
in the first premaxillary tooth of Sinornithosaurus' holotype. However,
Milner could have reversed the terms again and the distal carina may be the
serrated one. In any case, Deinonychus, Dromaeosaurus and probably
Utahraptor and Saurornitholestes differ in having both carinae
serrated on premaxillary teeth. Most of Sinornithosaurus' anterior teeth
are unserrated (as is reported for Microraptor), except for a few serrations
on the first one, as mentioned above. The tooth is said to be D-shaped, but
dromaeosaurines are often said to exhibit this condition as well, though it
is quite different from the tyrannosauroid morphology. This tooth has a few
strange characters as well- the enamel is striated, some serrations have a lingual
midline ridge, and others have been combined in labial view.
Comments- Discovered in 1852, Owen originally (1854) described this taxon
as a lizard, tentatively referring the tibia and fibula DORCM G 914 to Nuthetes
as well (later referred to the atoposaurid crocodilian Theriosuchus by
Seeley in 1893). Owen later (1879) identified Nuthetes as a crocodilian
and referred additional teeth to the taxon, as well as dermal armor called granicones.
Nuthetes was first identified as a dinosaur by Lydekker (1888), and as
a theropod by Zittel (1911), who placed it in the Coeluridae. Swinton (1934)
and Chakravarti (1935) referred it to the Megalosauridae. It was even synonymized
with Megalosaurus itself by Romer (1956). Delair (1959) believed the
granicones to be from a thyreophoran, and Galton (1986) referred them to Echinodon
(now recognized as a heterodontosaurid). They have been recently reidentified
as solemydid turtle limb and tail(?) scutes (Barrett et al., 2002). Though most
often viewed as a lizard or juvenile megalosauroid or carnosaur last century,
Nuthetes was reidentified as a juvenile dromaeosaurid by Milner (1999,
2002). Bonde (2012) thought three larger teeth (including NHMUK R 15870 and 15878)
were probably congeneric with Dromaeosauroides, while Nuthetes
itself could be "a more generalized tetanuran/neotheropod."
Milner assigns Nuthetes to the Dromaeosauridae based solely on the high
DSDI, though she also compares the Mackelian groove and posterior extent of
tooth tips to Deinonychus. She distinguishes it from "velociraptorines"
by the absence of apically inclined serrations, despite the fact she earlier
describes and illustrates these in the taxon. Although Milner says dromaeosaurines
are distinguished from Nuthetes by their low DSDI and that Nuthetes compares
most closely to "velociraptorines", she never actually assigns it
to the Velociraptorinae. No comparison or mention is made to non-dromaeosaurid
dromaeosaurs.
Other taxa besides "velociraptorines" have high DSDI's however. Within
coelurosaurs, these include basal tyrannosauroids (Guanlong, Dilong, Eotyrannus)
and Richardoestesia.
That Richardoestesia is not discussed by Milner is confusing, as she
lists it in her table of DSDI values as having a range almost identical to Nuthetes
(1.06-1.53). Richardoestesia gilmorei has a Mackelian groove that
is shallow like dromaeosaurids and an upper row of circular foramina. Further
details are difficult to make out, though an anterior foramen may be present.
It also has unfused interdental plates like Nuthetes. Notably, R.
gilmorei exhibits similar heterodonty, with some teeth being quite elongate
and others being shorter. And like dromaeosaurids, at least some teeth have
tips extending posterior to their distal base. Nuthetes falls within
the range of R. gilmorei for labiolingual thickness vs. FABL, height
vs. FABL. Serration morphology is similar, with blunt rounded tips slightly
hooked apically. Richardoestesia also has mesial serrations that vary
in extent down the carina, from extending the complete crown edge to being absent.
One of the characteristics of the genus is the minute distal serration size
(5-12/mm), and this agrees with Nuthetes (4.5-8/mm). The tentatively
identified premaxillary tooth of Richardoestesia is only serrated on
one carina, with a cross section similar to Dromaeosaurus. Two important
differences are that Nuthetes has interdenticle slits and has no significant
basal constriction on lateral teeth. Also, Nuthetes shows none of the
elongate straight teeth with convex distoapical margins known to exist in the
anterior dentary of Richardoestesia, nor the elongate "stacked banana"
serrations seen mesially in some teeth of that genus. Although R. isosceles
is more similar in that it lacks basal constriction, it is far less recurved
than Nuthetes, never as short as Nuthetes dentary teeth, has even
smaller interdenticle slits, and never has rounded or pointed serrations.
Eotyrannus differs in having fused interdental plates and larger serrations
(2.6/mm), though its Mackelian groove is shallow. Guanlong also has larger
serrations (3.8/mm) and is said to have labiolingually thick lateral teeth,
unlike Nuthetes. Dilong is more difficult to compare as it has
been only preliminarily described. Its dentary tooth apices do extend posterior
to the distal base, it has unfused interdental plates, its lateral teeth are
highly compressed labiolingually and lack basal constriction, heterodont in
the dentary at least, and have serrations which are rounded and sometimes asymmetrical.
The serrations are separated by interdenticle slits as in Nuthetes. The
premaxillary teeth are D-shaped, as described in Nuthetes, though whether
Nuthetes' premaxillary tooth is truly D-shaped is uncertain as noted
above. Unfortunately, the serration size, dentary foramina and medial dentary
morphology are still undescribed. Nuthetes cannot be distinguished from
Dilong from what is currently described of the latter, but this this
may change as more material is discovered of Nuthetes and more is described
on Dilong. It is provisionally placed by Dilong here.
References- Owen, 1854. On some fossil reptilian and mammalian remains
from the Purbecks. Quarterly Journal of the Geological Society of London. 10,
420-433.
Owen, 1961. Monograph on the fossil Reptilia of the Wealden and Purbeck formations.
Part V. Lacertilia (Nuthetes, etc.). [Purbeck]. The Palaeontological
Society, London. 1858, 31-39.
Owen, 1878. On the fossils called 'granicones'; being a contribution to the
histology of the exo-skeleton in 'Reptilia'. Journal of the Microscopical Society.
1, 233-236.
Owen, 1879. Monograph on the fossil Reptilia of the Wealden and Purbeck formations.
Supplement no. IX. Crocodilia (Goniopholis, Brachydectes, Nannosuchus,
Theriosuchus, and Nuthetes). The Palaeontographical Society. 1879,
1-19.
Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British
Museum (Natural History). Part I. Containing the Orders Ornithosauria, Crocodilia,
Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum (Natural
History), London. 1-309.
Seeley, 1893. On a reptilian tooth with two roots. Annals and Magazine of Natural
History, s 6. 12, 227-230.
Zittel, 1911. Grundz�ge der Pal�ontologie (Pal�ozoologie). II.
Abteilung. Vertebrata. Druck und Verlag von R. Oldenbourg, M�nchen. 1-598.
Swinton, 1934. The Dinosaurs. George Allen & Unwin, London. 233 pp.
Chakravarti, 1935. Is Lametasaurus indicus an armored dinosaur?. The
American Journal of Science, series 5. 30, 138-141.
Romer, 1956. Osteology of the Reptiles, University of Chicago Press. 772 pp.
Delair, 1959. The Mesozoic reptiles of Dorset. Proceedings of the Dorset Natural
History and Archaeology Society. 80, 52-90.
Galton, 1986. Herbivorous adaptations of Late Triassic and Early Jurassic dinosaurs.
In Padian (ed.). The Beginning of the Age of Dinosaurs. Cambridge University
Press. 203-221.
Milner, 1999. [title] Life and Environments in Purbeck Times. Abstracts, [pp].
Milner, 2002. Theropod dinosaurs of the Purbeck Limestone Group,
southern England. In Milner and Batten (eds.). Life and Environment in
Purbeck Times. Special Paper in Palaeontology. 68, 191-201.
Barrett, Clarke, Brinkman, Chapman and Ensom, 2002. Morphology, histology and
identification of the 'granicones' from the Purbeck Limestone Formation (Lower
Cretaceous: Berriasian) of Dorset, southern England. Cretaceous Research. 23,
279-295.
Bonde, 2012. Danish dinosaurs: A review. In Godefroit (ed.). Bernissart Dinosaurs
and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 434-451.
N. sp. (Mazin, Billo-Bruyat, Pouech and Hantzpergue, 2006)
Berriasian, Early Cretaceous
unnamed unit, Charente, France
Material- (CHEm03.537) tooth
References- Mazin, Billo-Bruyat, Pouech and Hantzpergue, 2006. The Purbeckian
site of Cherves-de-Cognac (Berriasian, Early Cretaceous, southwest France):
Acontinental ecosystem accumulated in an evaporitic littoral depositional environment.
9th International Symposium, Mesozoic Terrestrial Ecosystems and Biota, Abstracts
and Proceedings volume. 84-88 and 169.
Pouech, Mazin and Billon-Bruyat, 2006. Microvertebrate biodiversity from Cherves-deCognac
(Lower Cretaceous, Berriasian: Charente, France). 9th International Symposium,
Mesozoic Terrestrial Ecosystems and Biota, Abstracts and Proceedings volume.
173.
"Stokesosauridae" Wu, Shi, Dong, Carr, Yi and Xu, 2019
Comments- This family name was
used four times by Wu et al. (2019) but is not valid as it is not
"explicitly indicated as intentionally new" (ICZN Article 16.1) or "be
accompanied by citation of the name of the type genus" (Article
16.2). Wu et al. used it to include Stokesosaurus, Juratyrant and Eotyrannus.
Reference- Wu, Shi, Dong, Carr, Yi and Xu, 2019. A new tyrannosauroid from the
Upper Cretaceous of Shanxi, China. Cretaceos Research. Journal
Pre-proof DOI: 10.1016/j.cretres.2019.104357
Stokesosaurus Madsen, 1974
S. clevelandi Madsen, 1974
= Iliosuchus clevelandi (Madsen, 1974) Galton, 1976
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of Morrison Formation, Colorado(?), South Dakota, Utah,
US
Holotype- (UUVP 2938) (~1.8 m) ilium (220 mm)
Paratype- (UUVP 2320) ilium (~330 mm)
Referred- ?(BYUVP 4862) ischia (521, 570 mm) (Britt, 1991)
?(BYUVP 5073) distal caudal vertebra (65 mm) (Britt, 1991)
?(BYUVP 8908) distal caudal vertebra (66 mm) (Britt, 1991)
?(UUVP 2455) basoccipital, partial parasphenoid, basisphenoid (Chure and Madsen,
1998)
?(UUVP 11689) furcula (Chure and Madsen, 1996)
?(lost) ilium (~120 mm) (Foster and Chure, 2000)
maxilla, complete braincase (Loewen, Sertich and Irmis, 2012)
Diagnosis- (after Benson, 2008) swollen rim around articular surface
of pubic peduncle; median ridge thicker than in Stokeosaurus langhami
and extending almost to dorsal margin of blade.
Comments- The ilium described by Foster and Chure (2000) may be Aviatyrannis.
Benson (2008) noted it differed from Stokesosaurus in having a strictly
vertical median ridge and a blade with a lower profile, so referred it to Tyrannosauroidea
indet.. The premaxilla UUVP 2999 originally referred to Stokesosaurus
by Madsen (1974) was later referred to Tanycolagreus (Carpenter et al.,
2005), but Benson feels it is more likely a ceratosaur. Benson also referred
the caudal vertebrae described by Brit (1991) and the braincase UUVP 2455 described
by Chure and Madsen (1998) to Theropoda indet., though he noted the vertebrae
do not differ significantly from those of S. langhami. Curtice and Wilhite
(1996) noted that the middle caudal described by Britt (BYUVP 5103) had since
been reassigned by Britt to Ceratosaurus. Nesbitt et al. (2009) noted
a furcula described by Chure and Madsen (1996) as Theropoda indet. may be Stokesosaurus,
as it resembles tyrannosauroids in being U-shaped with expanded epicleideal
processes. Loewen et al. (2012) noted Cleveland-Lloyd quarry materials referrable
to Stokesosaurus including a premaxilla, maxilla, complete braincase,
two ilia and two ischia. The ilia are probably the holotype and paratype, but
the ischia may be new as BYUVP 4862 are from Dry Mesa. The premaxilla may be
UUVP 2999 which is from Cleveland-Lloyd, and the braincase could be UUVP 2455
from the Cleveland-Lloyd except that that specimen is not complete, preserving
only the posteroventral portion.
References- Madsen, 1974. A new theropod dinosaur from the Upper Jurassic
of Utah. Journal of Paleontology. 48, 27-31.
Galton, 1976. Iliosuchus, a Jurassic dinosaur from Oxfordshire and Utah.
Palaeontology 19, 587-589.
Chure and Madsen, 1996. On the presence of furculae in some non-maniraptoran
theropods. Journal of Vertebrate Paleontology. 16(3), 573-577.
Curtice and Wilhite, 1996. A re-evaluation of the Dry Mesa Dinosaur Quarry sauropod
fauna with a description of juvenile sauropod elements. In Huffman, Lund and
Godwin (eds.). Geology and Resources of the Paradox Basin. Utah Geological Association
Guidebook 25, 325-338.
Chure and Madsen, 1998. An unusual braincase (?Stokesosaurus clevelandi)
from the Cleveland-Lloyd Dinosaur Quarry, Utah (Morrison Formation; Late Jurassic).
Journal of Vertebrate Paleontology. 18(1), 115-125.
Foster and Chure, 2000. An ilium of a juvenile Stokesosaurus (Dinosauria,
Theropoda) from the Morrison Formation (Upper Jurassic: Kimmeridgian), Meade
County, South Dakota. Brigham Young University Geology Studies. 45, 5-10.
Carpenter, Miles and Cloward, 2005. New small theropod from the Upper Jurassic
Morrison Formation of Wyoming. In Carpenter (ed.). The Carnivorous Dinosaurs.
Indiana University Press. 23-48.
Benson, 2008. New information on Stokesosaurus, a tyrannosauroid (Dinosauria:
Theropoda) from North America and the United Kingdom. Journal of Vertebrate
Paleontology, 28(3), 732-750.
Nesbitt, Turner, Spaulding, Conrad and Norell, 2009. The theropod furcula. Journal
of Morphology. 270, 856-879.
Loewen, Sertich and Irmis, 2012. The early evolution of tyrannosauroid dinosaurs:
New anatomical, phylogenetic and biogeographic evidence. Journal of Vertebrate
Paleontology. Program and Abstracts 2012, 129.
Juratyrant
Brusatte and Benson, 2013
= "Notomegalosaurus" Carrano, 1998
= "Juratyrant" Brusatte and Benson, 2012 online
J. langhami (Benson, 2008) Brusatte and Benson, 2013
= "Juratyrant" langhami (Benson, 2008) Brusatte and Benson,
2012 online
= Stokesosaurus langhami Benson, 2008
Early Tithonian, Late Jurassic
Kimmeridge Clay, England
Holotype- (OUMNH J.3311) fourth or fifth cervical vertebra (56 mm), partial
first dorsal vertebra, mid dorsal vertebra (75 mm), partial mid dorsal vertebra
(77 mm), partial posterior dorsal vertebra (90 mm), partial posterior dorsal
vertebra (86 mm), incomplete sacrum (440 mm), partial proximal caudal vertebra
(74 mm), partial proximal caudal vertebra (82 mm), incomplete proximal caudal
vertebra (89 mm), proximal caudal vertebra (96 mm), distal caudal centrum, two
partial chevrons, four transverse processes, incomplete ilia (523 mm), pubes
(545 mm), incomplete ischia, incomplete femora (~667 mm), tibiae (one partial;
680 mm), fragment
Diagnosis- (after Benson, 2008) reduced and dorsally raised postzygapophyses
on last dorsal vertebra; prominent hyposphene of fifth sacral vertebra; median
ridge of the ilium is narrower and does not continue as far toward the perimeter
of the blade; swollen ridge is not present around the pubic peduncle; ischial
apron with ‘folded’ appearance; fibular flange continues as distinct
low ridge to proximal end of tibia.
Comments- The holotype was discovered in 1984, and was listed as the tetanurine "Notomegalosaurus" sp.
in Carrano's (1998) thesis. It was first published as "Undescribed theropod"
by Weishampel et al. (2004), with further details given in Martill et al. (2006)
and identified as a new species of Stokesosaurus by Benson (2007) in
an abstract. Benson (2008) described it the next year as such, but Brusatte
and Benson (2013) placed it in its own genus as the characters shared with S.
clevelandi were found to be more widely distributed and langhami
grouped with Eotyrannus in their tree. Though the online version of the
paper appeared in February 2012, it was only physically published in 2013.
References-
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004.
Dinosaur Distribution. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria
Second Edition. University of California Press. 517-606.
Martill, Naish and Earland, 2006. Dinosaurs in marine strata: Evidence from
the British Jurassic, including a review of the allochthonous vertebrate assemblage
from the marine Kimmeridge Clay Formation (Upper Jurassic) of Great Britain.
In Colectivo Arqueol�gico-Paleontol�gico Salense (ed.). Actas
de las III Jornadas Internacionales sobre Paleontolog�a de Dinosaurios
y su Entorno. Salas de los Infantes, Burgos. 47-83.
Benson, 2007. A new Jurassic tyrannosauroid from the Tithonian (Late Jurassic)
of Dorset, UK, representing a large-bodied species of the American genus Stokesosaurus.
Journal of Vertebrate Paleontology. 27(3), 47A.
Benson, 2008. New information on Stokesosaurus, a tyrannosauroid (Dinosauria:
Theropoda) from North America and the United Kingdom. Journal of Vertebrate
Paleontology, 28(3), 732-750.
Benson, 2009. The taxonomy, systematics and evolution of the British
theropod dinosaur Megalosaurus. PhD thesis, University of Cambridge. [pp]
Brusatte and Benson, 2012 online. The systematics of Late Jurassic tyrannosauroids
(Dinosauria: Theropoda) from Europe and North America. Acta Palaeontologica
Polonica. http://dx.doi.org/10.4202/app.2011.0141
Brusatte and Benson, 2013. The systematics of Late Jurassic tyrannosauroid theropods
from Europe and North America. Acta Palaeontologica Polonica. 58(1), 47-54.
Eotyrannus Hutt, Naish, Martill,
Barker and Newbery, 2001
= “Gavinosaurus” Kelly, 1998
= “Lengosaurus” Kelly, 1998
= “Kittysaurus” Hargreaves, 2001
= “Fusinasus” Hutt, 2002
E. lengi Hutt, Naish, Martill, Barker and Newbery 2001
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (IWCMS : 1997.550; = MIWG 1997.550) (~4.5 m; subadult) premaxilla,
partial maxilla, fused nasals (220 mm), incomplete lacrimal, incomplete quadrate,
partial dentaries, partial surangulars, several teeth, axial centrum, axial
neural arch, cervical neural arch, cervical centrum, mid dorsal centrum, three
incomplete dorsal centra (52 mm), several dorsal central fragments, partial
dorsal neural arch, several dorsal rib fragments, sacral centrum (71 mm), three
partial caudal centra, two partial distal caudal vertebrae, incomplete scapulae
(~280 mm), incomplete coracoid, incomplete humeri (235 mm), proximal radius,
partial ulna, distal carpal I, metacarpal I, phalanx I-1, manual ungual I (~115
mm on curve), proximal metacarpal II, phalanx II-1, phalanx II-2 (85 mm), manual
ungual II (~108 mm on curve), proximal metacarpal III, phalanx III-1, phalanx
III-3 (70 mm), ilial fragments, incomplete tibia, partial fibula, metatarsal
II (250 mm), distal metatarsals III, phalanx III-1 (87 mm), metatarsal IV (~260
mm), phalanx IV-3/4, pedal phalanges, pedal ungual, additional material
Referred- ?(Dinosaur Expeditions Centre coll.; Glyn's tibia) incomplete
tibia (Mattsson, pers. comm. 2015)
Comments- Eotyrannus was discovered in 1997 and was first announced
by Martill and Hutt in 1998 at the Dinosaur Society conference "British
Dinosaurs - Their Lifes and Times". A number of names were suggested in
popular press for the genus- "Gavinosaurus", "Lengosaurus"
and "Kittysaurus", though "Fusinasus" seems to have been
a serious consideration by the authors. After the preliminary description by
Hutt et al. (2001), it was monographed by Naish in his 2006 thesis. Naish and
Martill (2007) provided new cranial and skeletal reconstructions based on this.
The monograph has been revised by Naish and Cau, and is due for publication
soon.
Mattsson (pers. comm. 2015) informs me of an incomplete tibia nicknamed 'Glyn's
tibia' which is tentatively referred to Eotyrannus. The specimen is on display
at the Dinosaur Expeditions Centre and is being studied by Siebert.
This is traditionally placed as a non-tyrannosaurid tyrannosauroid as suggested
by nearly all analyses, though a few place it slightly closer to birds (e.g.
Lee and Worthy, 2011).
References- Hutt and Hutt, 1998. A new small theropod dinosaur from the
Isle of Wight. SVPCA 1998. [pp]
Kelly, 1998. Is this man our Indiana Jones? The Daily Mail. 10-7-1998.
Hutt, Naish, Martill, Barker and Newbery, 2001. A preliminary account of a new
tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern
England. Cretaceous Research. 22, 247-242.
Hargreaves, 2001. He's daddy of the dinosaurs. The News. May 10, 13.
Hutt, 2002. Mr Leng's dinosaur. The Geological Society of the Isle of Wight
Newsletter. 2(6), 12-14.
Naish, 2006. The osteology and affinities of Eotyrannus lengi and Lower
Cretaceous theropod dinosaurs from England. PhD thesis, University of Portsmouth.
[pp]
Naish and Martill, 2007. Dinosaurs of Great Britain and the role of the Geological
Society of London in their discovery: Basal Dinosauria and Saurischia. Journal
of the Geological Society. 164, 493-510.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Naish and Cau, in press.
Tyrannosauridae sensu Brochu, 2003
Definition- (Alectrosaurus olseni + Gorgosaurus libratus + Albertosaurus
sarcophagus + Daspletosaurus torosus + Alioramus remotus +
Tarbosaurus bataar + Tyrannosaurus rex)
Reference- Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from
a nearly complete skeleton and high-resolution computed tomographic analysis
of the skull. SVP Memior 7. 138 pp.
"Teihivenator" Yun,
2017
"T." macropus (Cope, 1868) Yun, 2017
= Laelaps macropus Cope, 1868
= Dryptosaurus macropus (Cope, 1868) Hay, 1902
Late Campanian-Early Maastrichtian, Late Cretaceous
Navesink Formation, New Jersey, US
Syntypes- (AMNH 2550) proximal tibia, distal tibia (95 mm wide)
?...(AMNH 2551) phalanx II-1 (93 mm), phalanges III-1 (83 mm)
?...(AMNH 2552) distal metatarsal II
?...(AMNH 2553) proximal metatarsal II or IV
Diagnosis- Indeterminate compared to Bistahieversor and Alioramus.
Other diagnoses- Previously (2010) this website proposed a diagnosis
for macropus combining "lateral tibial malleolus at same level as
medial malleolus" and "paired proximoventral processes on pedal phalanges
II-1 and III-2", but with the former now known in multiple tyrannosauroids
and the latter referrable to a different taxon, I agree with Brownstein that
the species is indeterminate though distinct from other named eastern American
tyrannosauroids.
In addition to those two, Yun (2017) listed more characters in his diagnosis.
Brownstein dismissed "medial tibial condyle is triangular, whereas lateral
tibial condyle is round" and "medial tibial condyle is positioned
higher than lateral tibial condyle" as being due to poor preservation and
erosion. Similarly, Brownstein interpreted the "small tubercle ... present
between the well-separated medial and lateral tibial condyle" to be merely
the less eroded portion of the bone surface. Brownstein noted that while the
"intercondylar notch is deep and “I” shaped" in AMNH 2550,
that Appalachiosaurus has a similar shape and that erosion could have
modified its depth. Finally, that the "cnemial crest can be seen at the
posterior view of [the] proximal tibia" is normal, and the opposite is
only present in Dryptosaurus due to erosion.
Comments- Leidy (1865) originally described this material as syntypes
of his new taxon Coelosaurus antiquus. Cope (1868) separated it as Laelaps
macropus, distinguishing it from Dryptosaurus (his Laelaps)
aquilunguis by its relatively longer pedal phalanges. He later (1870)
described it in more detail and illustrated some elements. Cope distinguished
it from "Coelosaurus" by its larger size and more expanded
distal tibia, and from Dryptosaurus by its anterior process on the lateral
tibial condyle. Matthew and Brown (1922) considered it probable that the specimen
was referrable to "Coelosaurus" antiquus after all, which has
been the consensus ever since on the rare times macropus is mentioned.
Gallagher (1997) photographed all the material, though note the distal tibial
piece is placed above the proximal piece in his figure. Holtz (2004) listed
it as an indeterminate tyrannosauroid without comment. Most recently, Yun was
mistakenly told the material was missing and published a short paper (Yun, 2017)
only using information from the literature (including this website, without
attribution) attempting to name the genus Teihivenator. However, the
syntypes are all accounted for and were being redescribed by Brownstein (2017)
as a chimaera of tyrannosauroid and ornithomimosaur elements. Notably, Yun's
paper is not available in physical form and lacks a ZooBank registration, making
the genus a nomen nudum (ICZN Article 8.5.3). While Yun did later register the
genus at ZooBank, Article 8.5.3 states that a name 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."
Since the work itself lacks such evidence, the name remains a numen nudum until
the publication of a corrigendum. Brownstein intends to designate a lectotype
in a future draft of his paper.
The tibia is tyrannosauroid based on the anterior process of the lateral tibial
condyle, and also matches tyrannosaurids more than ornithomimids in the rounded
posterolateral edge of the medial condyle. The lateral corner is placed more
posteriorly than Alectrosaurus, Appalachiosaurus or Tyrannosaurus,
but is more similar to Albertosaurus and Dryptosaurus. The more
open triangular posterior groove resembles Appalachiosaurus and Dryptosaurus
more than Alectrosaurus, Albertosaurus or Tyrannosaurus.
The distal tibia has a less ventrally projecting lateral malleolus than most
other tyrannosauroids except Bistahieversor and Alioramus. Brownstein
(2017) asssigned distal metatarsal AMNH 2552 and proximal metatarsal AMNH 2553
to either Tyrannosauroidea or Ornithomimosauria, noting their color and wear
differs from the tibia, but may indicate these two specimens were associated.
The three pedal phalanges were referred to Ornithomimosauria by Brownstein,
based in part on their paired ventral proximoventral projections. These speciemns
are retained under the macropus heading here, pending publication and
official lectotype designation by Brownstein.
References- Leidy, 1865. Memoir of the extinct reptiles of the Cretaceous
formations of the United States. Smithsonian Contributions to Knowledge. 14,
1-135.
Cope, 1868. On the genus Laelaps. American Journal of Science. 2(66),
415-417.
Cope, 1870. Synopsis of the extinct Batrachia, Reptilia and Aves of North America.
Transactions of the American Philosophical Society. 14, 1-252.
Hay, 1902. Bibliography and Catalogue of the Fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 1-868.
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.
Gallagher, 1997. When Dinosaurs Roamed New Jersey. 176 pp.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds). The
Dinosauria (second edition). University of California Press, Berkeley. 111-136.
Brownstein, 2017. Theropod specimens from the Navesink Formation and their implications
for the diversity and biogeography of ornithomimosaurs and tyrannosauroids on
Appalachia. PeerJ Preprints. 5:e3105v1.
Yun, 2017. Teihivenator gen. nov., a new generic name for the tyrannosauroid
dinosaur "Laelaps" macropus (Cope, 1868; preoccupied by Koch,
1836). Journal of Zoological and Bioscience Research. 4(2), 7-13.
Moros Zanno, Tucker, Canoville, Avrahami, Gates and Makovicky, 2019
M. intrepidus Zanno, Tucker, Canoville, Avrahami, Gates and Makovicky, 2019
Late Cenomanian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Holotype- (NCSM 33392)
(~50-100 kg; >7 year old subadult) incomplete femur (~355 mm), incomplete tibia (~440 mm), fragmentary metatarsal
II, incomplete metatarsal IV (~270 mm), phalanx IV-3 (27.55 mm), phalanx IV-4 (18.88 mm)
Referred- ?(CM 71399) premaxillary tooth (Fiorillo, 1999)
?(NCSM 33276; COI-10) premaxillary tooth (6 mm) (Avrahami, Gates, Heckert,Makovicky and Zanno, 2018)
?(NCSM 33393) premaxillary tooth (11.34x5.65x4.04 mm) (Zanno, Tucker, Canoville, Avrahami, Gates and Makovicky, 2019)
?(OMNH coll.) premaxillary tooth (~9 mm), five lateral teeth (~15.7-~31.5 mm) (Kirkland and Parrish, 1995)
Diagnosis- (after Zanno et al.,
2019) semicircular tuberosity on anteromedial femoral shaft originating
at distalmost extent of anterior trochanter; sinuous articular facet on
medial aspect of fourth metatarsal for contact with third metatarsal;
transversely compressed, subtriangular distal articular condyle of
fourth metatarsal in distal view; distal articular surface of fourth
metatarsal exhibiting hypertrophied anterolateral aspect, confluent
with a deeply incised, striated extensor groove that grades
indistinctly into the lateral collateral ligament pit
Comments- Kirkland and Parrish (1995) mentioned remains of "tyrannosaurids represented by an aublysodontid; cf. Alectrosaurus", listed as "an early tyrannosaurid" cf. Alectrosaurus
sp. by Kirkland et al. (1997) and Cifelli et al. (1999). The latter
wrote "each theropod taxon is represented by isolated teeth" and that
"the faunal list for the Mussentuchit local fauna is based on more than
5,000 catalogued specimens in the collection of the OMNH."
Regarding the Robison eggshell site, Fiorillo (1999) said "one small
tooth from this site (CM 71399) has both the anterior and posterior
carinae on the lingual side of the tooth. The top half of the tooth is
missing but the cross section of the tooth is decidedly
D-shaped." From the OMNH's microvertebrate project, Kirkland et
al. (2016) figure six "tyrannosaurioid teeth, smallest tooth is a
diagnostic premaxillary tooth." Avrahami (2018; published as Avrahami
et al., 2018) described a series of microvertebrates from the Cliffs of
Insanity locality including tyrannosauroid premaxillary tooth NCSM
33276 found on July 25 2015. Zanno et al. (2019) described
premaxillary tooth NCSM 33393 from the Suicide Hill locality as
Tyrannosauroidea indet. alongside NCSM 33276 and noted the former has
an autapomorphic "deep, obliquely oriented groove incising the lingual
ridge." This is lacking in NCSM 33276 and is undescribed in CM
71399 and not visible in the photographed OMNH premaxillary
tooth. Notably no Mussentuchit lateral teeth have been described
as tyrannosauroid, but those photographed by Kirkland et al. match NCSM
33268 (~20 mm) from the Cliffs of Insanity site and Morphotype 1 of
Frederickson et al. (2018) (5-35 mm) in size. Due to the presence
of the large basal coelurosaur Siats
in the same formation however, these are not referred to
Tyrannosauroidea here. Early referrals of these teeth to
Aublysodontidae and/or Alectrosaurus
were probably due to the widespread view in the 1990s that these were
primitive tyrannosauroids with more slender teeth than tyrannosaurids
and lacking serrations on their premaxillary teeth. The latter
feature has since been discovered to be variable in basal
tyrannosauroids (e.g. true in Yutyrannus and Xiongguanlong) and juvenile tyrannosaurids, while Alectrosaurus has no convincingly referred dental remains.
Moros is based on a hindlimb
discovered in 2012 (Johnson, 2019) and described by Zanno et al. (2019)
from the Stormy Theropod site. Mussentuchit tyrannosauroid teeth
are provisionally referred to the taxon here.
Zanno et al. (2019) recovered it as a non-eutyrannosaur more derived than Dilong
in Brusatte's TWiG matrix, a non-eutyrannosaur more derived than
"stokesosaurids" in Carr's tyrannosauroid analysis and a eutyrannosaur
sister to Appalachiosaurus, Alioramus and tyrannosaurids in Loewen's tyrannosauroid analysis.
References- 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.
Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial Cretaceous
vertebrates from the Cedar Mountain Formation, Emery County, Utah: The Mussentuchit
local fauna. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison eggshell
site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. In Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 259-268.
Kirkland, Suarez, Suarez and Hunt-Foster, 2016. The Lower Cretaceous in
east-central Utah - the Cedar Mountain Formation and its bounding
strata (field trip guide). Geology of the Intermountain West. 3,
101-228.
Avrahami, 2018. Paleobiodiversity of a new microvertebrate locality
from the Upper Cretaceous Mussentuchit Member, Cedar Mountain
Formation, Utah: Testing morphometric multivariate approaches for
quantifying shape variation in microvertebrate specimens. Masters
thesis, North Carolina State University. 181 pp.
Avrahami, Gates, Heckert,Makovicky and Zanno, 2018. A new
microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain
Formation: Insights into the paleobiodiversity and paleobiogeography of
early Late Cretaceous ecosystems in western North America. PeerJ.
6:e5883.
Frederickson, Engel and Cifelli, 2018. Niche partitioning in theropod
dinosaurs: Diet and habitat preference in predators from the uppermost
Cedar Mountain Formation (Utah, U.S.A.). Scientific Reports. 8:17872.
Johnson, 2019. New dinosaur T. rex relative Moros intrepidus discovered. The News & Observer. February 21, 2019.
Zanno, Tucker, Canoville, Avrahami, Gates and Makovicky, 2019.
Diminutive fleet-footed tyrannosauroid narrows the 70-million-year gap
in the North American fossil record. Communications Biology. 2:64.
Alectrosaurus Gilmore, 1933
A. olseni Gilmore, 1933
= Albertosaurus olseni (Gilmore, 1933) Paul, 1988
Middle-Late Campanian, Late Cretaceous
AMNH 136,
Iren Dabasu Formation, Inner Mongolia, China
Lectotype- (AMNH 6554) two incomplete manual unguals(?), pubic foot fragment, femur
(647 mm), tibia (732 mm), proximal fibula, astragalus (77 mm wide), calcaneum,
metatarsal I (~62.8 mm), phalanx I-1 (61.3 mm), pedal ungual I (43.4+ mm), metatarsal
II (460.7, 470.7 mm), phalanx II-1 (114 mm), phalanx II-2 (88.2 mm), pedal ungual
II (35.2+ mm), metatarsal III (486 mm), phalanx III-1 (109.5 mm), phalanx III-2
(83.2 mm), phalanx III-3 (67.5 mm), pedal ungual III (38 mm), metatarsal IV
(478.2 mm), phalanx IV-1 (~79.6 mm), phalanx IV-2 (~67 mm), phalanx IV-3 (52.5
mm), phalanx IV-4 (38.9 mm), pedal ungual IV, metatarsal V (109.9+ mm)
Middle-Late Campanian, Late Cretaceous
CCDP #9,
Iren Dabasu Formation, Inner Mongolia, China
Referred- ?(IVPP coll.) partial skeleton (Dong, Currie and Russell, 1989)
Middle-Late Campanian, Late Cretaceous
Erenhot,
Iren Dabasu Formation, Inner Mongolia, China
?(IVPP 170788104 or 180788-104) teeth (Currie, Rigby and Sloan, 1990)
Late Cretaceous
southeastern Mongolia
?(IGM coll.) several partial skeletons (Currie, 2001)
Diagnosis- (after Carr, 2005a) spike-like process extends from the caudodorsal
surface of the medial condyle of the femur; oval scar on the posterior surface
of the femur is lateral to the midline; medial margin of the joint surface for
the astragalus on the tibia is straight; shallow muscular fossa extends posteriorly
from the medial pocket of the fibula; abrupt expansion in length of the anterior
margin of the joint surface for the tibia on the fibula; tendon pit adjacent
to the ventrolateral buttress of the astragalus undercuts the medial surface
of the buttress; base of lateral flange of metatarsal I is triangular; metatarsal
I anteroposteriorly narrow; apex of distal joint surface of metatarsal I situated
medial to the midline of the bone; lateral collateral ligament pit of metatarsal
I does not extend anteroventrally adjacent to the distal joint surface; lateral
condyle of pedal phalanx I-1 extends above the dorsal surface of the bone; ventral
lateral condyle of pedal phalanx I-1 extends ventrolaterally; medial ligament
pit of pedal phalanx I-1 is small and circular; dorsolateral condyle of metatarsal
II is pediculate; medial edge of medial ventral condyle of metatarsal II extends
below the shaft surface; spur extends from the posterolateral edge of metatarsal
II above the distal joint surface; dorsal margin of proximal surface of pedal
phalanx II-2 is pointed; lateral dorsal condyle of pedal phalanx II-2 in dorsal
view reaches the midlength of the collateral ligament pit; deep and narrow cleft
separates distal condyles of pedal phalanx II-2; center of the flexor groove
of pedal phalanx II-2 is convex; flexor tubercle of pedal unguals II-IV are
hypertrophied and reach the level of the proximal joint surface; proximal joint
surface of pedal digits II-IV bear a low vertical ridge on the midline; dorsal
lateral and ventral lateral condyles of metatarsal III are pediculate; in anterior
view the dorsal margin of the distal condyle of metatarsal III is horizontally
oriented; the medial edge of the distal joint surface of metatarsal III extends
beyond the shaft margin; the supracondylar pit of metatarsal III is shallow;
in ventral view, the distal joint surface of metatarsal III is hyperextended
onto the shaft; shaft of metatarsal III elongate; pedal digit III is short;
in distal view the lateral condyle of pedal phalanx III-1 is significantly deeper
than the medial condyle; the distal joint surface of pedal phalanx III-1 is
deeply concave; in ventral view the posterior margin of the distal condyle of
pedal phalanx III-1 is convex; in distal view the distal condyles of pedal phalanx
III-2 are narrow and deep; in ventral view the lateral ridge that bounds the
flexor groove of pedal phalanx III-2 is a prominent keel; rugosities are absent
above the collateral ligament pits of pedal phalanx III-3; in dorsal view, the
wide posterior region of the shaft of pedal phalanx III-3 is limited to the
posterior third of the shaft; in medial view the scar posterodorsal to the collateral
ligament pit is low in pedal phalanx III-3; in dorsal view the dorsal ridge
of pedal ungual III does not follow the midline; the distal joint surface of
metatarsal IV is pediculate except for the medial ventral condyle; the lateral
distal condyle of metatarsal IV is hyperextended onto the ventral surface of
the bone; the cleft that separates the condyles of metatarsal IV extends onto
the distal end of the joint surface; in lateral view the distal margin of the
lateral distal condyle of pedal phalanx IV-1 is flattened; in proximal view
pedal phalanx IV-2 is narrow; in dorsal view the lateral condyle of pedal phalanx
IV-2 extends ventrolaterally; in dorsal view the joint surface of the lateral
distal condyle of pedal phalanx IV-3 extends proximally; a narrow cleft separates
the distal condyles of pedal phalanx IV-4; the medial collateral ligament pit
of pedal phalanx IV-4 is situated close to the dorsal margin of the bone; a
longitudinal groove excavates the distal third of the ventral surface of pedal
phalanx IV-4; the dorsal half of the joint surface for metatarsal IV on metatarsal
III is dilated anteriorly.
Comments-
The lectotype hindlimb AMNH 6554) was discovered on April 25 1923 at
Third Asiatic Expedition field site 136, while a partial forelimb (AMNH
6368) was found on May 4 at field site 138, 30 meters away.
Andrews (1932) first mention the former as "the complete hindlimb of a
large carnivorous dinosaur. The leg lay doubled up just as the
great reptile had died millions of years ago." Gilmore (1933)
made each a syntype of his new taxon of deinodontid, Alectrosaurus olseni,
noting "in the field they were thought to pertain to the same
individual." but that he preferred to treat them as two
individuals. They were considered the same taxon based on the
manual unguals (questionably associated in the case of AMNH 6554)
"being laterally compressed, strongly curved, and having sharply
pointed extremities", which are characters broadly true of almost all
theropod manual unguals. Barsbold (1976) was the first to
consider AMNH 6368 wrongly assigned, stating "As new materials from the
MPR* show, a large ungual phalanx previously attributed to the manus of
Alectrosaurus (Gilmore, 1933)
does not really belong to it" (translated), citing the still
undescribed IGM 100/50 from Bayanshiree which includes "a small ungual
phalanx of the first manual digit, quite typical for
tyrannosaurids." Further, he noted "A large, laterally compressed
ungual phalanx, similar in structure and form to that attributed to Alectrosaurus,
belongs to another previously unknown dinosaur (under study) found
there. This dinosaur does not belong to Tyrannosauridae.", which is a
reference to the also undescribed Segnosaurus.
In their redescription, Mader and Bradley (1989) describe AMNH 6368 in
detail and place it in Segnosauridae, and it has been viewed as
therizinosaurian since then. As explained by Mader and Bradley,
the type listing by White (1973) combined the syntype materials, so
that Welles and Long (1974) officially declared the hindlimb as the
lectotype when they stated "we here designate this specimen, AMNH 6554,
the type of the species." Zanno (2010: Fig. 9D) figured the
manual unguals of AMNH 6554 as therizinosaurian without comment,
although their non-tyrannosaurid characters could also be
plesiomorphically shared with e.g. Dryptosaurus.
Dong et al. (1989) first reported Aublysodon
from the July 1988 Sino-Canadian expedition (CCDP), and Currie et al.
(1990) stated "Identical teeth [to Dinosaur Park juvenile tyrannosaurid
'Aublysodon'] recently were recovered from the Iren Dabasu Formation at Erenhot, People's Republic of China (IVPP 170788104). The Asian "Aublysodon" teeth belong to Alectrosaurus
(Perle pers. comm. 1989..." Similarly, Dong (1992) reports "In
July 1988, the expedition of the CCDP came to Erenhot (Fig.85) where
they collected ... teeth of ... large theropods (tyrannosaurid)."
Dong et al. first reported that in July 1988 "A partial skeleton of Alectrosaurus was discovered too late to collect", and Dong (1993) followed that up by writing "An incomplete skeleton of Alectrosaurus
was found by Currie [in 1988], but was not excavated until the return
expedition of 1990", which was at CCDP site #9 based on Currie and
Eberth's (1993) table 3. They further noted "Perle (1977) ... has
been studying more recently discovered postcranial specimens (Perle,
pers. comm. 1989)" and that "The absence of denticles on the
premaxillary teeth (Perle, pers. comm. 1989; IVPP 180788-104) suggests
that it should be included in the Aublysodontinae." The
similar field numbers to Currie et al.'s suggest one is a typo, and
serrationless premaxillary teeth are a juvenile character of
tyrannosaurines but also known in some basal tyrannosauroids (Yutyrannus, Xiongguanlong). Currie (2001) reported "Several partial, undescribed skeletons of Alectrosaurus
collected from southeastern Mongolia are in the collections of the
museum in Ulaanbaatar, and another new specimen was recently collected
from Erenhot in China", the latter seemingly being the one mentioned by
Dong et al.. Finally, a fragmentary skull reported by Brochu
(2003) and under study by Carr (2005b) found in 1922 (AMNH 6266;
mistakenly called AMNH 6556 by Carr) may belong to Alectrosaurus (perhaps even to the type individual) and currently resolves close to Jinbeisaurus and Timurlengia. None of
these can be confirmed as belonging to the genus however, and may be juveniles
of a larger tyrannosaurid reported by Gilmore (1933) for instance.
Perle (1977) described two partial specimens (IGM 100/50, 100/51)
as Alectrosaurus, including skull material, which have formed much of
the basis for our understanding of the genus. However, Carr (2005a) found these
specimens differ from the holotype and could find no support for their referral.
Numerous specimens from Kazakhstan, Uzbekistan and Tajikistan have been referred
to Alectrosaurus, primarily due to Nessov (1995). None of these show
autapomorphies of Alectrosaurus, however (Carr, 2005a). Most are teeth
which cannot be compared to the taxon, though the myth of Cenomanian-Santonian
labiolingually narrow tyrannosauroid teeth being Alectrosaurus has spread.
The tooth reported by Gangloff (1998) from the Chandler Formation (Albian-Cenomanian)
of Alaska is actually from Dromaeosaurus (Fiorillo and Gangloff, 2000).
Carr (2005a, b) redescribes Alectrosaurus, noting an extremely large
number of hindlimb apomorphies which he interpreted as indicating enhanced cursorial
abilities. In a hindlimb-only phylogenetic analysis of tyrannosaurids, Alectrosaurus
was resolved as the sister taxon of Dryptosaurus. However, the topology
of tyrannosauroids is quite different from that in analyses based on cranial
characters. When added to Brusatte et al.'s 2010 tyrannosauroid analysis,
Alectrosaurus is more basal than other taxa included here in Tyrannosauroidea.
References- Andrews, 1932. The New Conquest of Central Asia. The American Museum of Natural History. 678 pp.
Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.
White, 1973. Catalogue of the genera of dinosaurs. Annals of Carnegie Museum.
44(9), 117-155.
Welles and Long, 1974. The tarsus of theropod dinosaurs: Annals of the South
African Museum. 44, 117-155.
Barsbold, 1976. New data on Therizinosaurus (Therizinosauridae, Theropoda). In Kramarenko, Luvsandansan, Voronin, Barsbold, Rozhdestvensky, Trofimov and Reshetov (Eds.).
Paleontology and Biostratigraphy of Mongolia. The Joint Soviet-Mongolian
Paleontological Expedition, Transactions. 3, 76-92.
Perle, 1977. On the first discovery of Alectrosaurus (Tyrannosauridae,
Theropoda) from the Late Cretaceous of Mongolia. Problemy Geologii Mongolii.
3, 104-113.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.
Mader and Bradley, 1989. A redescription and revised diagnosis of the syntypes
of the Mongolian tyrannosaur Alectrosaurus olseni. Journal of Vertebrate
Paleontology. 9(1), 1-55.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press. 107-125.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People’s Republic
of China. Cretaceous Research. 14, 127-144.
Dong, 1993. The field activities of the Sino-Canadian Dinosaur Project
in China, 1987-1990. Canadian Journal of Earth Sciences. 30(10),
1997-2001.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii. Institute for Scientific Research on the Earth's
Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation
(Cretaceous) of northern Alaska, with speculations on Arctic dinosaur paleoecology.
Journal of Vertebrate Paleontology. 20(4), 675-682.
Currie, 2001. Theropod dinosaurs from the Cretaceous of Mongolia. In Benton,
Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia.
434-455.
Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from a nearly
complete skeleton and high-resolution computed tomographic analysis of the skull.
Society of Vertebrate Paleontology Memior. 7, 138 pp.
Carr, 2005a. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with
special reference to North American forms. PhD thesis. University of Toronto.
1170 pp.
Carr, 2005b. A reappraisal of tyrannosauroids from Iren Dabasu, Inner Mongolia,
People's Republic of China. Journal of Vertebrate Paleontology. 25(3), 42A.
Zanno, 2010. A taxonomic and phylogenetic re-evaluation of
Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic
Palaeontology. 8(4), 503-543.
Suskityrannus Nesbitt, Denton, Loewen, Brusatte, Smith, Turner, Kirkland, McDonald and Wolfe, 2019
= "Zunityrannus"
Molina-Perez and Larramendi, 2019
S. hazelae Nesbitt, Denton, Loewen, Brusatte, Smith, Turner, Kirkland, McDonald and Wolfe, 2019
Middle Turonian, Late Cretaceous
Lower Moreno Hill Formation, New Mexico, US
Holotype-
(MSM P4754; Fred) (~2 m, ~10 kg, juvenile) incomplete premaxillae,
incomplete maxillae (98, 102 mm), jugal fragments, distal quadrate
fragment, parietal fragments, braincase fragments, partial vomer,
partial palatines, incomplete dentaries, partial splenials, posterior
mandible, anterior cervical vertebra with partial fused cervical ribs
(23 mm), mid cervical vertebra with partial fused cervical ribs (29
mm), anterior dorsal centrum (~20 mm), partial sacral centrum, distal
metatarsal II, distal metatarsal III, distal metatarsal IV, fragments
Paratype- (MSM P6178) (3+ year
old juvenile) partial postorbital, frontal, anterior dentary,
incomplete anterior cervical vertebra, mid dorsal vertebra (21 mm), mid
dorsal neural spine, two posterior dorsal centra, partial anterior
sacral centrum (17 mm), proximal caudal centrum (29 mm), four mid
caudal vertebrae (35 mm), three distal caudal vertebrae (41 mm), neural
arches, partial scapula, two partial manual unguals, incomplete pubes,
femora (one incomplete; ~230 mm), incomplete tibiae (~230, ~231 mm with
astragalus), partial fibulae, astragali (35 mm trans), incomplete
metatarsal II, phalanx II-1, incomplete metatarsal III, incomplete
metatarsal IV, phalanx IV-3, fragments
Diagnosis- (after Nesbitt et
al., 2019) distal condyles of femur strongly mediolaterally compressed
with crista tibiofibularis that merges smoothly into the lateral
condyle laterally; medial condyle of proximal end of tibia hooked
posteromedially.
Comments- The holotype was discovered in 1997, and reported by
Wolfe and Kirkland (1998) as a "new small dromaeosaurid theropod"
and a "small (dromaeosaurid?) theropod." The paratype was found
in 2000, including more postcranial elements.
Kirkland and Wolfe (2001) referred to the taxon as a basal coelurosaur, as did
Holtz in 2001 (DML). Indeed, casts of the skeleton are available to be purchased
as the "Zuni coelurosaur". Pringle (2001) photographed much of the
skeleton in her popular article, where the specmen was nicknamed Fred. In 2004,
Denton et al. referred to it as a basal tetanurine based on several characters
discussed below. More recently, McDonald et al. (2010) referred to it as "a
new taxon of small basal tyrannosauroid." The television program "Planet
Dinosaur" called the taxon "Zunityrannus" in episode 6, aired
in 2011. The companion book (BBC, 2011) seemingly uses Sinotyrannus due
to an editorial mistake, as in the closed captions for the show, so "Zunityrannus"
remained an unpublished name until listed as a nomen nudum by
Molina-Perez and Larramendi (2019). Nesbitt et al. (2019) named and officially described the taxon as Suskityrannus, recovering it as sister to Timurlegia, Xiongguanglong
and eutyrannosaurs in Brusatte's TWiG matrix and Brusatte and Carr's
tyrannosauroid matrix. They found it emerged as an alioramin
eutyrannosaur in Loewen et al.'s tyrannosauroid matrix.
References- Wolfe and Kirkland, 1998. Zuniceratops christopheri
n. gen. & n. sp., a ceratopsian dinosaur from the Moreno Hill Formation
(Cretaceous, Turonian) of west-central New Mexico. In Lucas, Kirkland and Estep
(eds.). New Mexico Museum of Natural History and Science Bulletin. 14, 307-317.
Holtz, DML 2001. https://web.archive.org/web/20201119060732/http://dml.cmnh.org/2001Jun/msg00634.html
Kirkland and Wolfe, 2001. First definitive therizinosaurid (Dinosauria; Theropoda)
from North America. Journal of Vertebrate Paleontology. 21(3), 410-414.
Pringle, 2001. The creature from the Zuni lagoon. Discover. August, 42-48.
Denton, Nesbitt, Wolfe and Holtz, 2004. A new small theropod dinosaur from the
Moreno Hill Formation (Turonian, Upper Cretaceous) of New Mexico. Journal of
Vertebrate Paleontology. 24(3), 302A.
Mcdonald, Wolfe and Kirkland, 2010. A new basal hadrosauroid (Dinosauria: Ornithopoda)
from the Turonian of New Mexico. Journal of Vertebrate Paleontology. 30(3),
799-812.
BBC, 2011. Planet Dinosaur. BBC, London. 240 pp.
Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288
pp.
Nesbitt, Denton, Loewen, Brusatte, Smith, Turner, Kirkland, McDonald
and Wolfe, 2019. A mid-Cretaceous tyrannosauroid and the origin of
North American end-Cretaceous dinosaur assemblages. Nature Ecology
& Evolution. 3, 892-899. DOI: 10.1038/s41559-019-0888-0
Embasaurus Riabinin, 1931
E. minax Riabinin, 1931
Berriasian-Hauterivian, Early Cretaceous
Neocomian Sands, Mount Koi-Kara, Kazakhstan
Syntypes- (subadult) partial ?posterior cervical centrum (~63 mm), mid
dorsal centrum (102 mm)
Diagnosis- (proposed) differs from Xiongguanlong in being over
170% larger and having less steeply angled cervical centra.
Previous diagnoses- While not providing a formal diagnosis, Riabinin
(1931) distinguished Embasaurus from Dryptosaurus because the
latter has shallower ventral concavities on its caudal centra than Embasaurus
does on its dorsal centrum (expected when comparing caudals to dorsals), from
Spinosaurus due to its non-opisthocoelous centra (a plesiomorphy), from
Ceratosaurus due to its supposedly more gently sloping ventral centrum
margin and flat articular surfaces (both known in Ceratosaurus), and
from Allosaurus for the same reasons plus the much shallower and less
angled ventral centrum margin of the dorsal (all of which are similar to mid
dorsals of Allosaurus).
Comments- The two syntype vertebral centra of Embasaurus were
discovered in 1927 and described by Riabinin in 1931.
The larger centrum is platycoelous or amphiplatyan, 107% longer than tall, 107%
wider than tall, has a ventral concavity 17% of centrum depth, and lacks pleurocoels
or a ventral keel. What may be the ventral part of the parapophysis is visible
anteriorly. The neural arch is unpreserved and was not fused to the centrum,
indicating the individual was not adult.
The smaller centrum is missing its anterior end, but its length can be estimated
using the posterior width and angle of anterior transverse expansion. While
it is much smaller (width ~51% of the larger centrum), this amount of disparity
is known between posterior dorsals and anterior cervicals in many theropods
(e.g. Majungasaurus). The angle between the posterior articular surface
and ventral edge suggests it is a cervical centrum, contra Riabinin. The posterior
surface is flat and 64% as wide as tall, with the length estimated at 94% of
the height. The ventral surface is keeled along its entire length and there
are no pleurocoels preserved, though they may have been present anteriorly.
Only the bases of the neural arches are preserved and the centrum interior is
hollow.
Riabinin (1931) referred Embasaurus to Carnosauria sensu lato based on
its size, and to Megalosauridae (also sensu lato, including allosauroids and
megalosauroids except Spinosaurus) based on "general form".
The few times authors have mentioned Embasaurus since (Nessov, 1995;
Currie, 2000) have repeated this possible identification without supporting
evidence. Molnar (1990) thought it was primitive due to the platycoelous dorsal
centrum and excluded it from Carnosauria sensu Gauthier (allosauroids and tyrannosaurids)
because of it, but most of the taxa he viewed as carnosaurs actually do have
roughly amphicoelous-amphiplatyan dorsals, besides the anterior dorsals of allosauroids.
Embasaurus can be excluded from Ceratosauria based on its lack of a posterior
cervical pleurocoel, though it is similar to many in having a ventral keel and
flat posterior surface. Megalosaurus itself and other megalosaurids differ
in having opisthocoelous cervical centra without a ventral keel, though the
posterior dorsal centra are roughly similar. Spinosaurids differ in the same
way. Some carcharodontosaurids have ventral keels (e.g. Carcharodontosaurus),
but all carnosaurs differ in having opisthocoelous cervical centra. Tyrannosauroids
are similar in being large and having non-opisthocoelous cervicals, and the
Early Cretaceous basal tyrannosauroid Xiongguanlong is similar in having
amphiplatyan cervicals with an elongate ventral keel on at least cervical ten
and lacking posterior dorsal pleurocoels. The only obvious difference is that
Embasaurus has a less steeply angled ventral edge on its cervical centrum
than any cervical of Xiongguanlong and that the subadult Embasaurus
individual was 170% larger than the adult Xiongguanlong holotype. Among
other tyrannosauroids, Dilong differs in having opisthocoelous cervicals
and amphicoelous dorsals that are more elongate, Stokesosaurus differs
in having platycoelous or opisthocoelous cervicals and amphicoelous dorsals,
and tyrannosaurids themselves lack ventral keels and have entirely pleurocoelous
dorsal centra. Therizinosaurs, ornithomimosaurs and oviraptorosaurs sometimes
get comparably large and have non-opisthocoelous cervicals, but the first two
have ventrally tranversely concave cervical centra, and all three have elongate
cervical vertebrae. Besides Xiongguanlong, the only close resemblence
is to basal tetanurines like Condorraptor and Szechuanosaurus? zigongensis,
which have platycoelous and keeled cervicals (in at least ~4 and ~10 in Condorraptor
and only in 9-10 in zigongensis) along with platycoelous posterior dorsals
that lack pleurocoels. These were generally extinct by the Cretaceous, though
specimens like Erectopus and the Baharija "Elaphrosaurus"
tibiae of Stromer may show they survived long enough for Embasaurus to
be a representative. Besides having posteriorly concave cervical centra, Condorraptor
differs in having its cervical keels only developed anteriorly as hypapophyses
and in lateral view the posteroventral centrum edge is convex in its cervicals.
Szechuanosaurus? zigongensis also differs in having posteriorly concave
cervical centra, with cervicals nine and ten broader and with more deeply concave
ventral edges. In conclusion, Embasaurus is most similar to the basal
tyrannosauroid Xiongguanlong, which is also close stratigraphically and
geographically. It is less similar to the generally earlier basal tetanurines,
so is referred here to Tyrannosauroidea. Within Tyrannosauroidea, it is more
derived than Dilong due to its short dorsal vertebrae but excluded from
Tyrannosauridae due to its lack of mid/posterior dorsal pleurocoels. As it differs
from all comparable taxa, it is not a nomen dubium as suggested by Molnar and
Holtz et al. (2004), neither of whom even compared it critically to other taxa.
References- Riabinin, 1931. Pozvonki dinozavra iz nizhnego mela Prikaspiyskikh
stepey. Zapiski Rossiyskogo Mineralogicheskogo Obshchestva. 60(2, number 1),
110-113.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California Press,
Berkeley, Los Angeles, Oxford. 306-317.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
Currie, 2000. Theropods from the Cretaceous of Mongolia. In Benton, Shishkin,
Unwin and Kurochkin (eds). The Age of Dinosaurs in Russia and Mongolia. Cambridge
University Press, Cambridge. 434-455.
Xiongguanlong Li, Norell,
Gao, Smith and Makovicky, 2010
= "Xiongguanlong" Li, Norell, Gao, Smith and Makovicky, 2009 online
X. baimoensis Li, Norell, Gao, Smith and Makovicky, 2010
= "Xiongguanlong baimoensis" Li, Norell, Gao, Smith and Makovicky,
2009 online
Late Aptian, Early Cretaceous
Middle Gray-variegated Beds, Xiagou Formation, Gansu, China
Holotype- (FRDC-GS JB16-2-1) (young adult; 272 kg) skull (504 mm), atlas,
axis (44 mm), third cervical vertebra (43 mm), fourth cervical vertebra (46
mm), fifth cervical vertebra (51 mm), sixth cervical vertebra (52 mm), seventh
cervical vertebra (60 mm), eighth cervical vertebra (60 mm), ninth cervical
vertebra (67 mm), tenth cervical vertebra (63 mm), first dorsal vertebra (62
mm), second dorsal vertebra (53 mm), third dorsal vertebra (58 mm), fourth dorsal
vertebra (55 mm), fifth dorsal vertebra (60 mm), sixth dorsal vertebra (58 mm),
seventh dorsal vertebra (56 mm), eighth dorsal vertebra, ninth dorsal vertebra
(55 mm), tenth dorsal vertebra (57 mm), eleventh dorsal vertebra, twelfth dorsal
vertebra, ilia (one incomplete, one fragmentary), femur (510 mm)
Diagnosis- (after Li et al., 2010) snout over two-thirds of skull length;
premaxillary teeth lack serrations (also in juvenile tyrannosaurines).
Other diagnoses- Li et al. (2010) also listed several symplesiomorphies
compared to tyrannosaurids (smooth nasal; lacrimal horn absent; quadrate not
pneumatic; single pair of cervical pleurocoels) and tyrannosauroid synapomorphies
absent in Dilong and Eotyrannus (premaxillary teeth with median
lingual ridge; enlarged nasal foramina absent; basicranium wider than long;
lateral processes on corners of axial neural spine).
Comments- Li et al.'s paper was first released electronically in April
2009 but not officially published until January 2010. The name was first published
in print in September 2009 as part of an SVP abstract (Makovicky et al., 2009),
but this was still a nomen nudum as abstracts are not allowed for official publication
(ICZN Article 9.10).
References- Makovicky, Li, Gao, Norell and Erickson, 2009. Two new coelurosaurs
from the Early Cretaceous Xinminpu Group of Gansu Province, China. Journal of
Vertebrate Paleontology. 29(3), 140A.
Li, Norell, Gao, Smith and Makovicky, 2010. A longirostrine tyrannosauroid from
the Early Cretaceous of China. Proceedings of the Royal Society B. 277(1679),
183-190.
You, Morschhauser, Li and Dodson, 2018. Introducing the Mazongshan
dinosaur fauna. Journal of Vertebrate Paleontology. 38(supp. 1), 1-11.
Jinbeisaurus Wu, Shi, Dong, Carr, Yi and Xu, 2019
J. wangi Wu, Shi, Dong, Carr, Yi and Xu, 2019
Late Cretaceous
Huiquanpu Formation, Shanxi, China
Holotype- (SMG V0003) (adult)
(skull ~660 mm) maxillae (one incomplete, one partial), incomplete
dentary, third cervical centrum, posterior cervical centrum, five
dorsal centra, distal pubis
Diagnosis- (after Wu et al.,
2019) interfenestral strut broad and covers more than 85% of the length
of the maxillary fenestra; ventral part of antorbital fossa, ventral to
the antorbital fenestra at the point just posterior to the tooth row,
is deeper than the subcutaneous surface below it; posteroventral end of
promaxillary recess on medial surface stops above fifth maxillary
alveolus; in medial view, the narrow septum between the promaxillary
recess and maxillary antrum becomes broad ventrally and bears a small
but deep fossa on its base; in medial view, the anterior part of the
maxillary antrum is nearly as large as the maxillary fenestra; dorsal
row of dentary foramina extends along the midheight of the dentary;
mesial and distal carinae bear a similar number of serrations per unit
length (about 16 at middle section of crown) in lateral teeth;
well-developed posterior pubic foot forms an acute angle (~70 degrees)
relative to the shaft.
Comments- This was discovered in 2008.
Wu et al. (2019) added Jinbeisaurus to Carr's tyrannosauroid analysis and recovered it in a polytomy with Timurlengia, AMNH 6556 and eutyrannosaurs.
Reference- Wu, Shi, Dong, Carr,
Yi and Xu, 2019. A new tyrannosauroid from the Upper Cretaceous of
Shanxi, China. Cretaceos Research. Journal Pre-proof DOI:
10.1016/j.cretres.2019.104357
Timurlengia Brusatte, Averianov,
Sues, Muir and Butler, 2016
T. euotica Brusatte, Averianov, Sues, Muir and Butler, 2016
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (ZIN PH 1146/16) (adult) incomplete braincase
Paratype- (ZIN PH 854/16) partial braincase
Referred- ?(CCMGE 432/12457) incomplete dorsal vertebra (93 mm) (Nessov,
1995)
?(CCMGE 433/12457-442/12457, except one) nine teeth (Nessov, 1995)
?(CCMGE 445/12457) pedal phalanx IV-1 (Nessov, 1995)
?(CCMGE 463/12457) pedal ungual (Nessov, 1995)
?(CCMGE 464/12457) pedal ungual (Nessov, 1995)
?(CCMGE 477/12475) distal caudal vertebra (Nessov, 1995)
?(CCMGE 485/12457) anterior lateral tooth (Nessov, 1995)
?(USNM 538123) (juvenile) dorsal neural arch (Averianov and Sues, 2012)
?(USNM 538131) partial posterior cervical vertebra (71 mm) (Averianov and Sues,
2012)
?(USNM 538132) (adult) anterior dorsal neural arch (Averianov and Sues, 2012)
?(USNM 538167) pedal ungual II (Averianov and Sues, 2012)
?(USNM 538181) manual ungual II (Averianov and Sues, 2012)
?(ZIN PH 2/16) maxillary fragment (Averianov and Sues, 2012)
?(ZIN PH 15/16) dentary fragment (Averianov and Sues, 2012)
?(ZIN PH 105/16) dorsal neural arch fragment (Averianov and Sues, 2012)
?(ZIN PH 106/16) dorsal neural arch fragment (Averianov and Sues, 2012)
?(ZIN PH 120/16) mid caudal vertebra (48 mm) (Averianov and Sues, 2012)
?(ZIN PH 121/16) astragalar fragment (Averianov and Sues, 2012)
?(ZIN PH 124/16) pedal ungual (58 mm) (Averianov and Sues, 2012)
?(ZIN PH 507/16) distal caudal vertebra (29.7 mm) (Averianov and Sues, 2012)
?(ZIN PH 619/16) manual ungual I (Averianov and Sues, 2012)
?(ZIN PH 671/16) anterior cervical centrum (79 mm) (Averianov and Sues, 2012)
?(ZIN PH 676/16) incomplete maxilla (261 mm) (Averianov and Sues, 2012)
?(ZIN PH 677/16) dentary fragment (Averianov and Sues, 2012)
?(ZIN PH 679/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 684/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 693/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 695/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 708/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 733/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 734/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 737/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 755/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 756/16) lateral tooth (Averianov and Sues, 2012)
?(ZIN PH 1033/16) premaxillary tooth (Averianov and Sues, 2012)
?(ZIN PH 1034/16) premaxillary tooth (Averianov and Sues, 2012)
?(ZIN PH 1035/16) premaxillary tooth (Averianov and Sues, 2012)
?(ZIN PH 1039/16) premaxillary tooth (Averianov and Sues, 2012)
?(ZIN PH 1239/16) (juvenile) posterior mandible (Averianov and Sues, 2012)
?(ZIN PH 1476/16) proximal caudal vertebra (80.4 mm) (Averianov and Sues, 2012)
?(ZIN PH 2296/16) distal quadrate (Averianov and Sues, 2012)
?(ZIN PH 2311/16) anterior dorsal centrum (Averianov and Sues, 2012)
?(ZIN PH 2312/16) anterior dorsal centrum (Averianov and Sues, 2012)
?(ZIN PH 2330/16) (juvenile) frontal (Averianov and Sues, 2012)
?(ZIN PH 2333/16) (juvenile) distal quadrate (Averianov and Sues, 2012)
?(ZIN PH 2350/16) (adult) posterior mandible (Averianov and Sues, 2012)
?(CCMGE 12457 and ZIN PH 16 coll.) several frontals, two posterior mandibles,
fifteen premaxillary teeth (5.3-19.2 mm), fifty-seven lateral teeth (to 65.2
mm), dorsal neural arch fragments, few caudal vertebrae (Nessov, 1995; Averianov
and Sues, 2012)
? distal femur, pedal phalanx II-1, distal pedal phalanx II-2 (Carr, 2005)
Diagnosis- (after Brusatte et al., 2016) diamond-shaped ventral projection
of supraoccipital that is excluded from foramen magnum; extremely short basal
tubera, approximately one third depth of occipital condyle; deep, funnel-like
otic recess that widely opens onto lateral surface of braincase and extends
far medially; large inner ear with robust semicircular canals.
Comments- The first Bissekty tyrannosauroid material was referred to
Allosaurus sp. by Sosedko (1937), then Deinodontidae by Efremov (1944).
Nessov (1995) referred material from the Bissekty Formation of Ukbekistan to
Alectrosaurus sp.. None exhibit Alectrosaurus synapomorphies and
several cannot be compared to the lectotype (Carr, 2005). This includes three
teeth (within CCMGE 433-442) among those later described by Averianov and Sues
(2012). He assigned CCMGE 445/12457 tentatively to juvenile Alectrosaurus
sp. as a metacarpal I. Carr thought it appeared to be a pedal phalanx IV-1,
but could not compare it in detail to confirm the taxonomic identification.
Nessov also assigned two pedal unguals (CCMGE 463/12457 and 464/12457) tentatively
to Alectrosaurus sp., but Carr could not compare them in detail to confirm
this identification. Additionally, thicker teeth (e.g. CCMGE 485/12457) were
assigned to Tyrannosauridae by Nessov, and premaxillary teeth were assigned
to Aublysodon sp., all of which Averianov and Sues include among the
tyrannosauroid teeth they describe. Nessov also assigned a distal caudal (CCMGE
477/12475) to Theropoda indet., which as reassigned to Tyrannosauroidea by Averianov
and Sues.
Material referred to Alectrosaurus by Ryan (1997) from the "Kulbecke
Formation" of Uzbekistan are actually from the Bissekty Formation (Nessov,
1995).
Archibald et al. (1998) reported tyrannosaurid teeth and bones from the Bissekty
Formation of Uzbekistan, three of which were examined by Carr (2005). He found
they were not referrable to Alectrosaurus, as they lack numerous apomorphies
of that genus when comparable.
Averianov (2007) notes 77 tyrannosaurid teeth are present, referring to the
77 lateral teeth later described by Averianov and Sues.
Averianov and Sues redescribed the Bissekty tyrannosauroid remains, beliving
them to pertain to one taxon due to the lack of variation and supposed lack
of other faunas with two tyrannosauroids (yet the Dinosaur Park Formation does,
so this is not valid). Coding them as one OTU, the material fell out more derived
than Raptorex and Dryptosaurus (based on the extensive frontal
supratemporal fossa, short cervical centra and rugose dorsal neural spines),
but less than Appalachiosaurus, Bistahieversor and Tyrannosauridae.
Brusatte et al. (2015) later proposed it was related to Xiongguanlong
based on small basal tubera and a diamond-shaped ventral supraoccipital process.
This was later officially published as Brusatte et al. (2016), who named the
taxon Timurlengia and described two new braincases as the holotype and
paratype. Using Brusatte's version of the TWG matrix, they found the holotype
to be in a polytomy with Juratyrant, Eotyannus and Dilong,
the non-braincase material to be sister to Xiongguanlong (thus more basal
than Averianov and Sues' analysis), and both combined to be between Dilong
and Xiongguanlong in a polytomy with Juratyrant and Eotyrannus.
References- Sosedko, 1937. Cemetery of vertebrates in the centre of Kyzyl-Kum
Desert. Sotsialisticheskaya Nauka i Tekhnika. 1937(5), 106-111.
Efremov, 1944. [Dinosaur horizon of Middle Asia and some questions of stratigraphy].
Izvestiya Akademii Nauk SSSR, Seriya Geologicheskaya. 3, 40-58.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii. Institute for Scientific Research on the Earth's
Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Ryan, 1997. Middle Asian dinosaurs. In Currie and Padian (eds.). Encyclopedia
of Dinosaurs. Academic Press. 442-444.
Archibald, Sues, Averianov, King, Ward, Tsaruk, Danilov, Rezvyi, Vereterunikov
and Khodjaev, 1998. Precis of the Cretaceous paleontology, biostratigtaphy and
sedimentology at Dzharakuduk (Turonian?-Santonian), Kyzylkum Desert, Uzbekistan.
Bulletin of the New Mexico Museum of Natural History and Science. 14, 21-27.
Sues and Averianov, 2004. Dinosaurs from the Upper Cretaceous (Turonian) of
Dzharakuduk, Kyzylkum Desert, Uzbekistan. Journal of Vertebrate Paleontology.
24(3), 51A-52A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Averianov and Sues, 2012. Skeletal remains of Tyrannosauroidea (Dinosauria:
Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan.
Cretaceous Research. 34, 284-297.
Brusatte, Carr, Averianov, Sues, Muir and Butler, 2015. Dinosaur dynasties:
Large theropod turnover in the Mid-Cretaceous as revealed by a new phylogeny
of tyrannosauroids and new fossils from Uzbekistan. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 98.
Brusatte, Averianov, Sues, Muir and Butler, 2016. New tyrannosaur from the mid-Cretaceous
of Uzbekistan clarifies evolution of giant body sizes and advanced senses in
tyrant dinosaurs. Proceedings of the National Academy of Sciences.
Brusatte, Muir, Averianov, Balanoff, Bever, Carr, Kundrat, Sues,
Williamson and Xu, 2016. Brains before brawn: Neurosensory evolution in
tyrannosauroid dinosaurs. Journal of Vertebrate Paleontology. Program
and Abstracts, 106.
undescribed tyrannosauroid (Granger and Berkey, 1922)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material-
(AMNH 6266; "AMNH 6556" of Carr, 2005) (small) incomplete lacrimal,
anterior jugal, quadratojugal, lateral ectopterygoid, posterior
pterygoid, premaxillary teeth, lateral teeth
Comments- Brochu (2003) noted "A box of bone fragments (AMNH 6266) from the same locality [as Alectrosaurus]
includes small tyrannosaurid skull bones (including a characteristic
jugal, lacrymal, quadratojugal, and D-shaped premaxillary tooth) that
might belong to the same individual. ... The skull parts are consistent
with Albertosaurus; for example, the jugal foramen is a dorsally-opening slit." He states it "had been originally catalogued as "Deinodon
sp." ..., but this was subsequently scratched off and "Theropoda
indet." written on in pencil", and the AMNH online catalogue does list
6266 as "Deinodon
? sp." Carr (2005) later reported "an undescribed, but shattered,
tyrannosauroid skull (AMNH 6556) from the same general area - Iren
Dabasu -" as the Alectrosaurus
lectotype, but as they are from different locations "there is no
evidence they are from the same individual." Carr states "The
presence of a secondary fossa in the antorbital fossa of the jugal
indicates the specimen is referable to Tyrannosauridae. The cornual
process of the lacrimal is similar to some juvenile tyrannosaurids in
that it is a low, laterally extending ridge. The lateral teeth are as
finely denticulate as tyrannosauroid teeth of the same basal crown
length from the Turonian of Uzbekistan." This is the "Iren Dabasu
taxon" in Carr's
tyrannosauroid analyses as of Carr et al. (2017), under study by Carr
and recovered in a
polytomy with Timurlengia, Jinbeisaurus
and eutyrannosaurs as of 2019. While this is certainly the same
specimen based on material preserved, the AMNH online catalog lists
this specimen number as being a saurischian metatarsal II with a
locality "8 mi. E. of station" which would place it among Third Asiatic
Expedition field sites 140-149 (while Alectrosaurus
is from 136 less than a mile south of the station). AMNH 6556 is
listed in the catalogue as collected on April 30, which matches Carr's
statement the skull was found in late April five days apart from Alectrosaurus'
lectotype (which was found on April 25). However, Mehling (pers.
comm. 6-2022) indicates AMNH 6556 is actually a metatarsal II and that
AMNH 6266 was discovered in 1922, so that Carr apparently got the
specimen number wrong and incorrectly used the metatarsal's discovery
date for the skull. The early discovery makes sense considering
the low specimen number and allows us to equate the material with
"portions of a small carnivorous dinosaur skull with two or three
teeth" found in the 1922 expedition as reported by Granger and Berkey
(1922) along with ornithomimid remains that are near certainly AMNH
6267-6268. If it was recovered with the latter specimens, AMNH
6266 would have been found between April 25 and May 7 at one of the
western AMNH quarries (131-138), and thus may be from the same locality
the Alectrosaurus type as
stated by Brochu. Alas, the only recorded locality information in
the AMNH card catalogue is Iren Dabasu (Mehling, pers. comm. 6-2022).
References-
Granger and Berkey, 1922. Discovery of Cretaceous and older Tertiary strata in Mongolia. American Museum Novitates. 42, 7 pp.
Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from a nearly
complete skeleton and high-resolution computed tomographic analysis of the skull.
Society of Vertebrate Paleontology Memior. 7, 138 pp.
Carr, 2005. A reappraisal of tyrannosauroids from Iren Dabasu, Inner Mongolia,
People's Republic of China. Journal of Vertebrate Paleontology. 25(3), 42A.
Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017. A new tyrannosaur
with evidence for anagenesis and crocodile-like facial sensory system.
Scientific Reports. 7:44942.
Eutyrannosauria Delcourt and Nelson Grillo, 2018
Definition- (Dryptosaurus aquilunguis + Tyrannosaurus rex) (modified from Delcourt and Nelson Grillo, 2018)
Reference- Delcourt and Nelson Grillo, 2018. Tyrannosauroids from the Southern
Hemisphere: Implications for biogeography, evolution, and taxonomy.
Palaeogeography, Palaeoclimatology, Palaeoecology. 511, 379-387.
DOI:10.1016/j.palaeo.2018.09.003
Dryptosauridae Marsh, 1890
Diagnosis- (after Brownstein,
2021) arctometatarsus in which metatarsal III lacks prominent
diaphysial bulge that articulates with II and IV; articular surface for
metatarsal V on metatarsal IV extends distally past proximal expansion;
ventral margin of metatarsal IV flat to concave in side view; no large
concavity along the ventral margin of metatarsal IV just proximal to
distal condyles; loss of groove or pit on lateral surface of distal end
of metatarsal IV.
Comments- Marsh (1890) proposed
the family Dryptosauridae for Cretaceous theropods (excluding
ornithomimids, which he assigned to Ornithopoda). The supposed
diagnostic characters are symplesiomorphic for theropods (limb bones
hollow; feet digitigrade; prehensile pedal digits) except for the very
small forelimbs. When the family was used in the 1900s it was
usually to emphasize the uniqueness of Dryptosaurus
among theropods (e.g. Paul, 1988; Carpenter et al., 1997) rather than
connect it with other genera. Brownstein (2021) proposed
inclusion of "Cryptotyrannus" in Dryptosauridae with a list of
synapomorphies.
References-
Marsh, 1890. Additional characters of the Ceratopsidae, with notice of new Cretaceous
dinosaurs. American Journal of Science. 39, 418-426.
Paul, 1988. Predatory Dinosaurs of the World. New York.
464 pp.
Carpenter, Russell, Baird and Denton, 1997. Redescription of the holotype of
Dryptosaurus aquilunguis (Dinosauria: Theropoda) from the Upper Cretaceous
of New Jersey. Journal of Vertebrate Paleontology. 17(3), 561-573.
Brownstein, 2021. Dinosaurs from the Santonian-Campanian Atlantic
coastline substantiate phylogenetic signatures of vicariance in
Cretaceous North America. Royal Society Open Science. 8, 210127.
Dryptosaurus Marsh, 1877
= Laelaps Cope, 1866 (preoccupied Koch, 1836)
Comments- While initially named Laelaps by Cope (1866), this genus
was preoccupied by a laelapid mite (Koch, 1836). Marsh (1877) renamed it Dryptosaurus.
Brusatte (2013) notes a revision of Late Cretaceous eastern North American tyrannosauroid
material is in progress, which should illuminate the identity of the non-holotype
specimens listed here.
References- Koch, 1836. Deutschlands Crustaceen, Myriapoden und Arachniden:
Ein beitrag zur deutschen, fauna, Volume 1. Pustet, Regensburg. 40 pp.
Cope, 1866. [On the remains of a gigantic extinct dinosaur, from the Cretaceous
Green Sand of New Jersey]. Proceedings of the Academy of Natural Sciences of
Philadelphia. 18, 275-279.
Marsh, 1877. Notice of a new and gigantic dinosaur. American Journal of Science
and Arts. 14, 87-88.
Brusatte, 2013. The phylogeny of basal coelurosaurian theropods (Archosauria:
Dinosauria) and patterns of morphological evolution during the dinosaur-bird
transition. PhD thesis, Columbia University. 944 pp.
D. aquilunguis (Cope, 1866) Marsh, 1877
= Laelaps aquilunguis Cope, 1866
= Megalosaurus aquilunguis (Cope, 1866) Osborn, 1898
Middle Maastrichtian, Late Cretaceous
Upper New Egypt Formation, New Jersey, US
Holotype- (ANSP 9995) (6.4 m; ~750 kg; adult) maxillary fragment, dentary
fragment, surangular fragment, several teeth, incomplete mid caudal centrum,
incomplete mid caudal centrum (115 mm), partial mid caudal vertebra (115 mm),
incomplete distal caudal vertebra (118 mm), incomplete distal caudal vertebra
(118 mm), incomplete distal caudal vertebra (113 mm), incomplete distal caudal
vertebra (108 mm), incomplete distal caudal vertebra (104 mm), incomplete distal
caudal vertebra (72 mm), two partial distal caudal vertebrae, incomplete humeri
(~300 mm), phalanx I-1 (~160 mm), manual ungual I or II (176 mm straight), phalanx
II-2 (126 mm), phalanx ?-? (48 mm; lost), incomplete pubes, partial ischium,
femur (781 mm), tibia (759 mm), partial fibula, partial astragalus (161 mm wide),
partial metatarsal III
....(AMNH 2438) metatarsal IV (396 mm)
Diagnosis- (after Carpenter et al., 1997) interdenticle spaces on maxillary
teeth half the width of denticles; flexor tubercle of manual ungual I begins
at proximal edge of articular surface and has minimal ventral projection.
(after Carr, 2005) medioventral heel of metatarsal IV absent.
(after Brusatte et al., 2011) combination of a reduced humerus (humerofemoral
ratio = 0.375) and large hand (phalanx I-1: femur ratio = 0.200); extremely
mediolaterally expanded ischial tubercle, ~1.7 times as wide as the shaft immediately
distally; ovoid fossa on medial surface of femoral shaft immediately proximal
to medial condyle, demarcated anteriorly by the mesiodistal crest and medially
by a novel crest; proximomedially trending ridge on anterior fibular surface
immediately proximal to iliofibularis tubercle; lip on lateral surface of lateral
condyle of astragalus prominent and overlapping the proximal surface of the
calcaneum; metatarsal IV with flat shaft proximally, resulting in a semiovoid
cross section that is much wider mediolaterally than deep anteroposteriorly.
Comments- The holotype was discovered in 1866. The sacral vertebrae originally
referred to the holotype, and used to suggest the specimen is immature, are
actually protostegid dorsal centra (Cope, 1872). Baird (1979) made them the
holotype of Pneumatoarthrus peloreus (see entry). The caudal vertebrae
have closed neurocentral sutures, suggesting an adult (Carpenter et al., 1997).
Four chevrons, a scapula and supposed sternum were noted/illustrated by Cope,
but are lost. The limb elements of AMNH 2438 are from the holotype locality,
and were thought by Huene (1932) to belong to the same specimen. They are preserved
differently than the holotype though, adding doubt to this assessment.
Long placed in its own family in an uncertain position among large theropods,
Baird and Horner (1979) placed it in the Tyrannosauridae based on the tentatively
referred femora. Paul (1988) felt it resembled coelurosaurs the most, and Denton
(1990) assigned it to that clade. Holtz (1996) found it to be a basal coelurosaur
as well, next to Deltadromeus and Bagaraatan, but more basal than
tyrannosaurids, Compsognathus, Ornitholestes and maniraptoriforms. Carpenter
et al. (1997) redescribed the material, noting resemblences to Betasuchus,
doubting its tyrannosaurid affinity, but ultimately preferring to keep it as
Theropoda incertae sedis. Denton (in Carpenter et al., 1997) however, was still
of the opinion Dryptosaurus was a coelurosaur, perhaps the most basal
form in that clade. It came out basal to all coelurosaurs except Proceratosaurus
in Holtz's (2000) analysis, and as a basal tyrannosauroid (by Stokesosaurus,
less derived than Eotyrannus and tyrannosaurids) in his unpublished 2001
analysis. Williamson and Carr (2001), Carr and Williamson (2002), Carr (2005)
and Brusatte et al. (2010) found it to come out as a tyrannosauroid more basal
than tyrannosaurids, Appalachiosaurus and Bistahieversor in their
analysis. Brusatte et al. (2011) redescriobed the material in depth.
References- Cope, 1866. [On the remains of a gigantic extinct dinosaur,
from the Cretaceous Green Sand of New Jersey]. Proceedings of the Academy of
Natural Sciences of Philadelphia. 18, 275-279.
Cope, 1868. On the genus Laelaps. American Journal of Science. 2(66),
415-417.
Cope, 1870. Some remains of a new Cretaceous turtle and on Laelaps. American
Philosophical Society Proceedings. 11, 515.
Cope, 1872. A description of the genus Protostega, a form of extinct Testudinata.
Proceedings of the American Philosophical Society. 12, 422-433.
Marsh, 1877. Notice of a new and gigantic dinosaur. American Journal of Science
and Arts. 14, 87-88.
Osborn, 1898. Paleontological problems. Science. 2, 145-147.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie, serie 1. 4(1-2), 1-361.
Baird, 1979. Pneumatoarthrus Cope, 1870, not a dinosaur but a sea-turtle.
Proceedings of the Academy of Natural Sciences of Philadelphia. 129, 71-81.
Paul, 1988. Predatory Dinosaurs of the World. New York.
464 pp.
Denton, 1990. A revision of the theropod Dryptosaurus (Laelaps)
aquilunguis (Cope 1869). Journal of Vertebrate Paleontology. 9(3), 20A.
Holtz, 1996. Phylogenetic analysis of the nonavian tetanurine dinosaurs (Saurischia:
Theropoda). Journal of Vertebrate Paleontology. 16(3), 42A.
Carpenter, Russell, Baird and Denton, 1997. Redescription of the holotype of
Dryptosaurus aquilunguis (Dinosauria: Theropoda) from the Upper Cretaceous
of New Jersey. Journal of Vertebrate Paleontology. 17(3), 561-573.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. Gaia. 15, 5-61.
Holtz, 2001. Pedigree of the tyrant kings: New information on the origin and
evolution of the Tyrannosauridae. Journal of Vertebrate Paleontology. 21(3),
62A-63A.
Williamson and Carr, 2001. Dispersal of pachycephalosaurs and tyrannosauroids
between Asia and North America. Journal of Vertebrate Paleontology. 21(3), 114A.
Carr and Williamson, 2002. Evolution of basal Tyrannosauroidea from North America.
Journal of Vertebrate Paleontology. 22(3), 41A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Brusatte, Benson and Norell, 2011. The anatomy of Dryptosaurus aquilunguis
(Dinosauria: Theropoda) and a review of its tyrannosauroid affinities. American
Museum Novitates. 3717, 53 pp.
D? sp. indet. (Casanova, 1987)
Campanian, Late Cretaceous
Blufftown Formation, Georgia
Material- (~7.1 m; ~1 ton) metatarsal II (~440 mm)
Comments- This cannot be compared to the holotype, and could be Appalachiosaurus
or another large theropod.
Reference- Casanova, 1987. Dryptosaurus sp., family Tyrannosauridae
a carnosaur, reported from Georgia. Fossils Quarterly. 6(3-4), 27-29.
D? sp. indet. (Gallagher, Paris and Spamer, 1986)
Late Campanian, Late Cretaceous
Marshalltown Formation, New Jersey, US
Material- (NJSM 12436) tooth (Gallagher, 1993)
(NJSM 13087) long bone shaft (Gallagher, 1993)
(NJSM 13095) tooth (Gallagher, 1993)
(NJSM 13096) (Gallagher, 1993)
(NJSM 13724) tooth (Gallagher, 1993)
(NJSM 14158) tooth (Gallagher, 1993)
(NJSM 14236) teeth, limb elements, phalanges (Gallagher, Paris and Spamer, 1986)
(NJSM 14404) tooth (Gallagher, 1993)
(NJSM 14434) (Gallagher, 1993)
(NJSM 14504) (Gallagher, 1993)
(NJSM 14682) proximal manual phalanx (Gallagher, 1993)
(NJSM 16664; cast) tooth (Gallagher, 1993)
Comments- Most of this material was called cf. Dryptosaurus by Gallagher
(1993), but may be Appalachiosaurus or another large theropod. NJSM 14664
was referred to Dryptosauridae indet. by Gallagher (1990).
References- Gallagher, Parris and Spamer, 1986. Paleontology, biostratigraphy,
and depositional environments of the Cretaceous-Tertiary transition in the New
Jersey Coastal Plain. The Mosasaur. 3, 1-36.
Gallagher, 1990. Dinosaurs-creatures of time. New Jersey State Museum Bulletin.
24, 43 pp.
Gallagher, 1993. The Cretaceous/Tertiary mass extinction event in the northern
Atlantic Coastal Plain. The Mosasaur. 5, 75-154.
D? sp. indet. (Horner, 1979)
Late Cretaceous
Marl Pits of James King, North Carolina, US
Material- (USNM 7189) two femora
Comments- Horner (1979) referred these to Dryptosaurus,
though this is not confirmed, as they are also similar to tyrannosaurids.
Reference- Horner, 1979. Upper Cretaceous dinosaurs from the
Bearpaw Shale (marine) of south-central Montana with a checklist of Upper Cretaceous
dinosaur remains from marine sediments of North America. Journal of Paleontology.
53, 566-577.
"Cryptotyrannus" Brownstein, 2021
Late Santonian-Early Campanian, Late Cretaceous
Merchantville Formation, Delaware, US
Material- (YPM VPPU.021795; intended holotype) (adult or subadult) incomplete metatarsal II (~400 mm), incomplete metatarsal
IV (~350 mm) (Horner, 1979)
?...(YPM VPPU.022416; intended paratype) partial mid caudal vertebra (Baird, 1986)
Diagnosis- (after Brownstein, 2021) differs from Dryptosaurus in- more gracile metatarsal IV (also in Moros and Alectrosaurus); triangular proximal outline of metatarsal IV; rectangular and mediolaterally compressed distal metatarsal IV (also in Moros); flexor sulcus of metatarsal IV deep;
Comments- Discovered in 1975, Horner (1979) initially listed
YPM VPPU.021795 as Ornithomimidae indet.. Baird (1986) mentioned
as "probably ornithomimid" "a partial metatarsus (PU 21795, R. Johnson
and R. Meyer coll.) and an anterior caudal centrum (PU 22416, W.
Cokeley coll.)". The latter was donated to the YPM in January
1979. Holtz (1992) mentions YPM PU 21795 as a referred "Coelosaurus" specimen,
noting it was subarctometatarsalian, more robust than ornithomimids, and had
only a slight ridge on metatarsal IV to back metatarsal III. He states it does
not resemble Dryptosaurus
and regards it as a distinct taxon, though he refrains from naming
it. Gallagher (1993) listed the metatarsals and all previously
identified eastern North American ornithomimosaur materials as juvenile
Dryptosaurus.
Brownstein (2017) described the metatarsals in detail in a preprint
which never led to a published paper in that journal. He
concluded it was "a new morphotype and possibly unnamed taxon of
tyrannosauroid" that could be differentiated from Dryptosaurus, Appalachiosaurus
and "Teihivenator", but ended up calling it Tyrannosauroidea
indet.. Dalman et al. (2017) independently described the
metatarsals and vertebra, also as Tyrannosauroidea indet., despite
similarly finding "they can be differentiated from other known
tyrannosauroids by the relatively straight proximal articular surface
of metatarsal II" which they believed "most likely represents a basal
condition shared with the Late Jurassic coelurid theropods."
Brownstein (2021) described the material again, this time as
"Dryptosauridae gen. et sp. nov.?", and seemingly intended to name the
taxon in an earlier draft as the name Cryptotyrannus
overlaps "Merchantville Taxon" in his figure 9 and the material is
listed and discussed as a holotype and paratype. However, he
ended up deciding to "take a conservative approach by suggesting novel
apomorphies for the Merchantville tyrannosauroid and thus presenting
Dryptosauridae as a multi-species clade while not erecting a new
name." While not addressed by prior workers, Brownstein (2021)
stated "The caudal vertebra was found close to the metatarsals at the
same locality, and because it belongs to a large theropod of the same
size as the one represented by metatarsals and shows closely comparable
preservation, it most likely belongs to the same individual."
References- Horner, 1979. Upper Cretaceous dinosaurs from the Bearpaw Shale (marine) of
south-central Montana with a checklist of Upper Cretaceous dinosaur
remains from marine sediments in North America. Journal of Paleontology.
53(3), 566-577.
Baird, 1986. Upper Cretaceous reptiles from the Severn Formation of Maryland.
The Mosasaur. 3, 63-85.
Holtz, 1992. An unusual structure of the metatarsus of Theropoda
(Archosauria: Dinosauria: Saurischia) of the Cretaceous. PhD Thesis, Yale University.
347 pp.
Gallagher, 1993. The Cretaceous/Tertiary mass extinction event in the northern Atlantic coastal plain. The Mosasaur. 5, 75-154.
Brownstein, 2017. A tyrannosauroid metatarsus from the Merchantville
Formation of New Jersey increases the diversity of non-tyrannosaurid tyrannosauroids
on Appalachia. PeerJ Preprints. 5:e3097v3.
Dalman, Jasinski and Lucas, 2017. First occurence of a tyrannosauroid
dinosaur from the Lower Campanian Merchantville Formation of Delaware,
USA. Memoir of the Fukui Prefectural Dinosaur Museum. 16, 29-38.
Brownstein, 2020. Reevaluation of dinosaur material from the Atlantic
coastal plain illuminates a bizarre new assemblage. The Society of
Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 88.
Brownstein, 2021. Dinosaurs from the Santonian-Campanian Atlantic
coastline substantiate phylogenetic signatures of vicariance in
Cretaceous North America. Royal Society Open Science. 8, 210127.
possible Dryptosauridae indet. (Gallagher, 1993)
Late Campanian, Late Cretaceous
Mount Laurel Formation, New Jersey, US
Material- (NJSM 14256) (~6-8 m) partial tooth (~25x8.99x4.90 mm)
Comments- Gallagher (1993) lists this specimen as "Cf. Dryptosaurus;
tooth, Mt. Laurel Fm., Hop Brook, Holmdel, Monmouth Co., N.J.".
Brownstein (2019) described the tooth in detail as Tyrannosauroidea
indet., recovering it sister to Tyrannosaurus
in Hendrickx' dental analysis. He further noted "NJSM GP 14256 is
narrow (CBW/CBL ~0.54) and possesses a lens-shaped basal cross-section,
indicative that it came from a tyrannosauroid outside Tyrannosauridae".
that it is specifically similar to Dryptosaurus
"in its dimensions, curvature and enamel crenulations" and thus given
its "very close spatio-temporal proximity to the holotype of Dryptosaurus, I suggest the tooth belongs to a closely related form."
References-
Gallagher, 1993. The Cretaceous/Tertiary mass extinction event in the northern Atlantic coastal plain. The Mosasaur. 5, 75-154.
Brownstein, 2019 New records of theropods from the latest Cretaceous of
New Jersey and the Maastrichtian Appalachian fauna. Royal Society Open
Science. 6, 191206.
undescribed possible dryptosaurid (Sereno, online 2001)
Early Cretaceous
Camp II, Mazongshan, Inner Mongolia, China
Material- (LH coll.) manual ungual (127 mm), manual elements
Comments-
This undescribed material was discovered by Sereno's team during
Dinosaur Expedition 2001. Sereno states the photographed ungual "and
the other bones of the hand we exposed clearly did not belong to the
diminutive forelimb of a tyrannosaur" and that "the bones of this new
predator were extremely fragile" so that further analysis requires
opening and prepapering the field jacket. The manual ungual
resembles Dryptosaurus most,
differing only in the more extensive area under the lateral groove proximally
and possibly the more anteriorly placed flexor tubercle (assuming the area isn't
broken). Perhaps the two are related.
Reference- Sereno, online 2001. https://web.archive.org/web/20020603195302/http://www.projectexploration.org/mongolia/u61001.htm
Raptorex Sereno, Tan, Brusatte,
Kriegstein, Zhao and Cloward, 2009
R. kriegsteini Sereno, Tan, Brusatte, Kriegstein, Zhao and Cloward,
2009
Late Valanginian-Hauterivian, Early Cretaceous?
Lujiatun Beds of the Yixian Formation?, Liaoning or Inner Mongolia, China
Holotype- (LH PV18) (2-3 year old juvenile) (~2.5 m, ~65 kg) incomplete
skull (~300 mm), incomplete mandibles, atlas (8 mm), axis (27 mm), third cervical
vertebra (28 mm), fourth cervical vertebra (29 mm), fifth cervical vertebra
(34 mm), sixth cervical vertebra (36 mm), seventh cervical vertebra (36 mm),
eighth cervical vertebra (34 mm), ninth cervical vertebra (35 mm), tenth cervical
vertebra (32 mm), three partial cervical ribs, first dorsal vertebra (28 mm),
second dorsal vertebra (30 mm), third dorsal vertebra (31 mm), fourth dorsal
vertebra (32 mm), fifth dorsal vertebra (32 mm), sixth dorsal vertebra (33 mm),
seventh dorsal vertebra (34 mm), eighth dorsal vertebra (36 mm), ninth dorsal
vertebra (36 mm), tenth dorsal vertebra (38 mm), eleventh dorsal vertebra (41
mm), twelfth dorsal vertebra (44 mm), thirteenth dorsal vertebra (45 mm), eighteen
partial to complete dorsal ribs, fused gastralium, five partial gastralia, sacrum
(47,47,48,49,46 mm), first caudal vertebra (40 mm), second caudal vertebra (40
mm), third caudal vertebra (41 mm), fourth caudal vertebra (41 mm), fifth caudal
vertebra (42 mm), sixth caudal vertebra (42 mm), seventh caudal vertebra (42
mm), eighth caudal vertebra (43 mm), ninth caudal vertebra (43 mm), tenth caudal
vertebra (43 mm), eleventh caudal vertebra (42 mm), scapula (151 mm), coracoid
(42 mm), humeri (99 mm), radius (52 mm), ulna (57 mm), metacarpal I (15 mm),
phalanx I-1 (26 mm), manual ungual I (~18 mm), phalanx II-1 (13 mm), phalanx
II-2 (~22 mm), incomplete ilia (335 mm), partial pubes (~289 mm), proximal ischia
(~225 mm), femora (338 mm), tibiae (one proximal; 397 mm), partial fibula, astragalus
(50 mm wide), pedal ungual I (17 mm), metatarsal II (245 mm), phalanx II-1 (55
mm), phalanx II-2 (35 mm), distal metatarsal III, phalanx III-1 (~62 mm), phalanges
III-2 (36, 37 mm), metatarsal IV (266 mm), pedal unguals IV (27, 27 mm)
Diagnosis- (after Sereno et al., 2009) narrow accessory pneumatic fossa
in antorbital fossa dorsal to maxillary fenestra; jugal suborbital ramus shallow
(transverse width approximately 60% vertical depth); vertical crest on iliac
blade dorsal to acetabulum absent (also in some Alioramus).
Comments- The holotype was collected privately without locality data,
so the exact provenence is unknown. Sereno et al. (2009) list measurements for
two left pedal phalanges III-1, one of which is much shorter and close to the
right III-2 in length so is probably III-2 instead. As the specimen is young,
it may be placed too basally in analyses like Sereno et al.'s and Brusatte et
al. (2010) which do not take into account ontogenetic changes in morphology
(Fowler et al., 2011; Tsuihiji et al., 2011). Fowler et al. (2011) also demonstrate
histology indicates Raptorex was more probably a 2-3 year old juvenile
than a 5-6 year old subadult. Brusatte (2013) states a restudy of Raptorex
and comparison to juvenile Tarbosaurus is in preparation by Sereno and
himself.
References- Sereno, Tan, Brusatte, Kriegstein, Zhao and Cloward, 2009.
Tyrannosaurid skeletal design first evolved at small body size. Science. 326(5951),
418-422.
Watanabe and Sereno, 2009. The forelimb of a new Tyrannosauridae (Dinosauria:
Theropoda) from Mongolia and its implications for forelimb evolution in tyrannosaurids.
Journal of Vertebrate Paleontology. 29(3), 198A.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Fowler, Woodward, Freedman, Larson and Horner, 2011. Reanalysis of "Raptorex
kriegsteini": A juvenile tyrannosaurid dinosaur from Mongolia. PLoS
ONE. 6(6), e21376.
Tsuihiji, Watabe, Tsogtbaatar, Tsubamoto, Barsbold, Suzuki, Lee, Ridgely, Kawahara
and Witmer, 2011. Cranial osteology of a juvenile specimen of Tarbosaurus
bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous)
of Bugin Tsav, Mongolia. Journal of Vertebrate Paleontology. 31(3), 497-517.
Brusatte, 2013. The phylogeny of basal coelurosaurian theropods (Archosauria:
Dinosauria) and patterns of morphological evolution during the dinosaur-bird
transition. PhD thesis, Columbia University. 944 pp.
Tyrannosauroidea incertae sedis
undescribed possible Tyrannosauroidea (Bakker, 2000)
Tithonian, Late Jurassic
Dinosaur National Monument, Brushy Basin Member of Morrison Formation, Utah, US
Material- (DNM uncatalogued) (Ford, pers. comm.)
Tithonian, Late Jurassic
Ward Facies, Brushy Basin Member of Morrison Formation, Wyoming, US
(WDIS 539) premaxillary tooth (~12 mm) (Bakker, 2000)
Comments-
Bakker (2000) figured WDIS 539 as "Dromaeosaurid, WDIS 539, anterior
premaxillary tooth, crown height restored 12 mm. Ward Facies. This
specimen is the earliest known dromaeosaurid." In an unpublished
appendix to Ford and Chure (2001), they list "Tyrannosaurid gen sp
indet DNM uncatalogued Dinosaur National Monument, Uintah County, Utah.
Morrison Formation, lower Bushy Basin Member Kimmeridgian, Late
Jurassic Valid." This material may be Stokesosaurus and/or Tanycolagreus.
References- Bakker, 2000 (as 1998). Brontosaur killers: Late Jurassic allosaurids as sabre-tooth cat
analogues. Gaia. 15, 145-158.
Ford and Chure, 2001. Ghost lineages and the paleogeographic and temporal distribution
of tyrannosaurids. Journal of Vertebrate Paleontology. 21(3), 50A-51A.
undescribed possible tyrannosauroid (Britt, Stadtman and Scheetz, 1996)
Barremian, Early Cretaceous
Yellow Cat Member of Cedar Mountain Formation, Utah, US
Material- teeth
Comments- This is noted as a possible tyrannosaurid.
Reference- Britt, Stadtman and Scheetz, 1996. The Early Cretaceous Dalton
Wells dinosaur fauna and the earliest North American titanosaurid sauropod.
Journal of Vertebrate Paleontology. 16(3), 24A.
unnamed tyrannosauroid (Gignac, Makovicky, Erickson and Walsh, 2010)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Wyoming, US
Material- (FMNH PR 2750) premaxillary tooth (~9 mm)
References- Gignac, Makovicky, Erickson and Walsh, 2010. A description of Deinonychus antirrhopus
bite marks and estimates of bite force using tooth indentation
simulations. Journal of Vertebrate Paleontology. 30(4), 1169-1177.
Zanno and Makovicky, 2011. On the earliest record of Cretaceous
tyrannosauroids in Western North America: Implications for an Early Cretaceous
Laurasian interchange event. Historical Biology. 23(4), 317-325.
undescribed possible tyrannosauroid (Thurmond 1974)
Aptian-Albian, Early Cretaceous
Middle Paluxy Formation of the Trinity Group, Texas, US
Material- (SMUSMP 62271) teeth
Reference- Thurmond, 1974. Lower vertebrate faunas of the Trinity Division
in north-central Texas. Geoscience and Man. 8, 103-129.
undescribed possible tyrannosauroid (Thurmond 1974)
Aptian-Albian, Early Cretaceous
Travis Peak Formation of the Trinity Group, Texas, US
Material- teeth
Reference- Thurmond, 1974. Lower vertebrate faunas of the Trinity Division
in north-central Texas. Geoscience and Man. 8, 103-129.
undescribed tyrannosauroid (Ullmann, Varricchio, Knell and Lacovara,
2010)
Albian, Early Cretaceous
Vaughn Member of the Blackleaf Formation, Montana, US
Reference- Ullmann, Varricchio, Knell and Lacovara, 2010. Taphonomy and
taxonomy of a vertebrate microsite in the Cretaceous Blacklaef Formation in
Southwest Montana. Journal of Vertebrate Paleontology. Program and Abstracts
2010, 179A.
unnamed tyrannosauroid (Bonde, 2008)
Albian, Early Cretaceous
Willow Tank Formation, Nevada, US
Material- premaxillary tooth (7x3x4 mm)
Reference- Bonde, 2008. Paleoecology and taphonomy of the Willow Tank
Formation (Albian), southern Nevada. Masters thesis, Montana State University.
96 pp.
undescribed Tyrannosauroidea (Krumenacker and Scofield, 2015)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2251/49858; Morph 1) lateral tooth (26.9x9.5x? mm) (Krumenacker
et al., 2016)
(IMNH 2251/49870; Morph 1) lateral tooth (31.4x11.1x7.1 mm) (Krumenacker et
al., 2016)
(IMNH 2251/49888; Morph 1) partial lateral tooth (Krumenacker et al., 2016)
(IMNH 2251/50088; Morph 1) lateral tooth (?x13.3x6.9 mm) (Krumenacker et al.,
2016)
(IMNH 2251/50089; Morph 1) lateral tooth (?x8.3x? mm) (Krumenacker et al., 2016)
(IMNH 2251/50848; Morph 1) lateral tooth (37x11.4x? mm) (Krumenacker et al.,
2016)
(IMNH 2251/50875; Morph 1) lateral tooth (16.8x7.8x? mm) (Krumenacker et al.,
2016)
(IMNH 2267/50105; Morph 3) lateral tooth (?x4.3x3.1 mm) (Krumenacker et al.,
2016)
(IMNH 2267/50106; Morph 3) lateral tooth (?x4.2x3.1 mm) (Krumenacker et al.,
2016)
(IMNH 2267/50107; Morph 3) lateral tooth (?x3.9x3 mm) (Krumenacker et al., 2016)
(IMNH 2267/50108; Morph 2) premaxillary tooth (7.3x3x2.2 mm) (Krumenacker et
al., 2016)
References- Krumenacker and Scofield, 2015. A diverse theropod tooth
assemblage from the Mid-Cretaceous (Albian-Cenomanian) Wayan Formation of Idaho.
Journal of Vertebrate Paleontology. Program and Abstracts 2015, 158.
Krumenacker, Simon, Scofield and Varricchio, 2016. Theropod dinosaurs from the
Albian-Cenomanian Wayan Formation of eastern Idaho. Historical Biology. DOI:
10.1080/08912963.2015.1137913
undescribed Tyrannosauroidea (Kirkland, Britt, Burge, Carpenter, Cifelli,
DeCourten, Eaton, Hasiotis and Lawton, 1997)
Late Cenomanian, Late Cretaceous
Dakota Formation, Utah, US
Material- (OMNH 24436) teeth
(juvenile) teeth (Kirkland et al., 1998)
? pubes, partial metatarsals (Kirkland, online 2007)
Comments- Kirkland et al. (1997) listed Tyrannosauridae indet. teeth,
while Kirkland et al. (1998) listed both Aublysodontinae indet. and Tyrannosaurinae
indet., implying both juvenile and adult individuals are preserved. Kirkland
(online) mentions and illustrates a pair of pubes he tentatively assigns to
a basal tyrannosaurid. He also mentions partial metatarsals of similar size,
which are provisionally listed here as they are too large to belong to other
known Dakota theropods (dromaeosaurids, troodontids, Richardoestesia
or Paronychodon).
References- 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.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau.
in Lucas, Kirkland and Estep (eds.). New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Kirkland, online 2007. https://scientists.dmns.org/sites/kencarpenter/ [offline]
undescribed Tyrannosauroidea (Kirkland, Lucas and Estep 1998)
Middle-Late Turonian, Late Cretaceous
Smoky Hollow Member of Straight Cliffs Formation, Utah, US
Material- (OMNH 21518) (juvenile) tooth (Parrish, 1999)
(OMNH 21524) (juvenile) tooth (Parrish, 1999)
(OMNH 24125) tooth (Parrish, 1999)
(OMNH 24436) tooth (Parrish, 1999)
Comments- Parrish (1999) listed OMNH 24125 and 24436 as Tyrannosauridae
and 21518 and 21524 as cf. Aublysodon.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. In Lucas, Kirkland and Estep (eds.). New Mexico Museum
of Natural History and Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian)
of southern Utah. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
undescribed Tyrannosauroidea (Kirkland, Lucas and Estep, 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Material- partial pes (Eaton pers. comm. 1996 to Kirkland, Lucas and Estep,
1998)
teeth? (Kirkland et al., 1998)
? (juvenile) material (Eaton et al., 1999)
? material (Eaton et al., 1999)
Comments- Kirkland et al. (1998) list both indet. Aublysodontinae and
indet. Tyrannosaurinae from this formation. Eaton et al. (1999) listed ?Tyrannosauridae
indet. and ?Aublysodon sp. from a Santonian possible Straight Cliffs
(or Wahweap?) locality.
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the
Colorado Plateau. In Lucas, Kirkland and Estep (eds.). New Mexico Museum of
Natural History and Science Bulletin. 14, 79-89.
Eaton, Diem, Archibald, Schierup and Munk, 1999. Vertebrate paleontology of
the Upper Cretaceous rocks of the Markagunt Plateau, southwestern Utah. In Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 323-333.
unnamed Tyrannosauroidea (Schwimmer et al., 1993)
Late Santonian-Middle Campanian, Late Cretaceous
Blufftown Formation, Alabama, Georgia, US
Material- (CCK-83-3-7) metatarsal shaft fragment
(CCK-84-4-7) partial radius
(CCK-84-4-8) partial ulna
(CCK-85-1-2) metatarsal shaft fragment
(CCK-87-5-1) incomplete metatarsal IV
(CCK-90-1-2) fragmentary pedal(?) phalanx
(CCK-90-5-1) metatarsal shaft fragment
(CCK-90-5-2) metatarsal shaft fragment
Comments- These may belong to the contemporaneous Appalachiosaurus.
Reference- Schwimmer, Williams, Dobie and Siesser, 1993. Late Cretaceous
dinosaurs from the Blufftown Formation in western Georgia and eastern Alabama.
Journal of Paleontology. 67(2), 288-296.
undescribed tyrannosauroid (Langston, 1960)
Early Campanian, Late Cretaceous
Mooreville Chalk Member of Selma Formation, Alabama, US
Material- (FMNH P27398) pedal phalanx
Reference- Langston, 1960. The vertebrate fauna of the Selma Formation
of Alabama. Part VI: The dinosaurs. Fieldiana. 3, 313-361.
unnamed Tyrannosauroidea (Cope, 1869)
Early Campanian, Late Cretaceous
Marl Pits of James King, Tar Heel Formation of the Black Creek Group, North Carolina, US
Material- (USNM 7189 in part; syntype of Hypsibema crassicauda) femoral shaft fragment (lost), distal femur (Cope, 1869)
Early Campanian, Late Cretaceous
Phoebus Landing, Tar Heel Formation of the Black Creek Group, North Carolina, US
(ANSP 15319) pedal phalanx III-3 (62 mm) (Miller, 1967)
(ANSP 15330) distal femur (Horner, 1979)
(ANSP 15332; 'ANSP 15331' of Miller, 1967) two incomplete teeth (FABL ~12.1, ~12.8 mm) (Miller, 1967)
(USNM 7199) tooth (20 mm) (Stephenson, 1912)
Comments-
Stephenson (1912) noted "Among the vertebrate remains from [Phoebus
landing] which are now in the U.S. National Museum, the following forms
have been identified by C. W. Gilmore:" "Carnivorous dinosaur
(Zatomus?)." Miller (1967) referred this to "Gorgosaurus?",
stating it is "flattened laterally, has a serrate anterior keel that
trends lingually toward the base of the crown, and a serrate, vertical,
posterior keel" and further states that "ANSP 15331 consists of an
identical, but incomplete tooth." He says the latter "is
identical to a tooth in the left jaw of Gorgosaurus libratus (USNM
12814). It matches the eleventh tooth, counted caudally, in size and
form." Baird and Horner (1979) note 'ANSP 15331' is actually
15332 and consists of two teeth, which they figure. Being an
earlier work, the authors write "They are as similar to Dryptosaurus
aquilunguis (Cope) as to Gorgosaurus (i.e. Albertosaurus)" and thus
"show no characteristics that would permit generic
identification."
ANSP 15319 was originally described by Miller (1967), who compared
it to pedal phalanx III-1 of Struthiomimus. Baird and Horner (1979) realized
it was a more distal phalanx (based on the ginglymoid proximal articular surface)
and referred it to cf. Ornithomimus as phalanx III-2. They stated
it closely resembled other ornithomimids, but cited Dryptosaurus? macropus
(AMNH 2551) as an example, while it's really a tyrannosauroid. In actuality,
ANSP 15319 differs from pedal phalanx III-2 of ornithomimids in being less elongate
and from III-2 in tyrannosauroids in being less transversely flared proximally
and distally. It is however, almost indistinguishable from pedal phalanx III-3
in both clades. It is provisionally referred to Tyrannosauridae here due to
size, as it is 24% larger than the largest Gallimimus specimen, but comparable
to a subadult Gorgosaurus.
Horner (1979) lists ANSP 15330 as Carnosauria indet., while it was described in depth and figured by Baird and Horner (1979).
Note anterior maxilla ANSP 15303 from Phoebus Landing has been reidentified as Deinosuchus.
References-
Stephenson, 1912. The Cretaceous formations. In Clark, Miller,
Stephenson, Johnson and Parker. The Coastal Plain of North Carolina.
North Carolina Geological and Economic Survey. Volume 3, 73-171.
Miller, 1967. Cretaceous vertebrates from Phoebus Landing, North
Carolina. Proceedings of the Academy of Natural Sciences of
Philadelphia. 119, 219-235.
Baird and Horner, 1979. Cretaceous dinosaurs of North Carolina. Brimleyana.
2, 1-28.
Horner, 1979. Upper Cretaceous dinosaurs from the
Bearpaw Shale (marine) of south-central Montana with a checklist of Upper Cretaceous
dinosaur remains from marine sediments of North America. Journal of Paleontology.
53(3), 566-577.
undescribed Tyrannosauroidea (Lubbe, Richter and Knotschke, 2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry, Germany
Material- ?(DFMMh/FV 382) tooth (9.7x5.8x3.4 mm) (Lubbe, Richter and Kn�tschke,
2009)
?(DFMMh/FV 790.5) (juvenile?) tooth (2.8x1.9x1.3 mm) (Lubbe, Richter and Kn�tschke,
2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry and/or Hannover, Germany
Material- (DFMMh and/or NLMH coll.) teeth
Comments- While seven teeth assigned to Velociraptorinae by Lubbe et
al. (2009), Gerke and Wings (2014) found four of these were Neotheropoda indet.,
megalosaurid and tyrannosauroid. Based on the information in Lubbe et al., FV
382 and 790.5 may be tyrannosauroid (less recurved and thicker labiolingually).
References- Lubbe, Richter and Kn�tschke, 2009. Velociraptorine
dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany. Acta Palaeontologica
Polonica. 54(3), 401-408.
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 Tyrannosauroidea (Lanser and Heimhofer, 2015)
Late Barremian-Early Aptian, Early Cretaceous
Balve-Beckum quarry, Germany
Material- (LWL MN Ba 15) premaxillary tooth (?x9.3x7 mm)
(LWL MN Ba 21) premaxillary tooth fragment
Reference- Lanser and Heimhofer, 2015. Evidence of theropod dinosaurs
from a Lower Cretaceous karst filling in the northern Sauerland (Rhenish Massif,
Germany). Pal�ontologische Zeitschrift. 89(1), 79-94.
undescribed Tyrannosauroidea (Efremov, 1944)
Late Cretaceous
Kshi-Kalkan, Almaty, Kazakhstan
Reference- Efremov, 1944. [Dinosaur horizon of Middle Asia and some questions
of stratigraphy]. Izvestiya Akademii Nauk SSSR, Seriya Geologicheskaya. 3, 40-58.
undescribed Tyrannosauroidea (Kordikova et al., 1996)
Turonian, Late Cretaceous
Kankazgan Formation, Kazakhstan
Reference- Kordikova, Gunnell, Polly and Kovrizhnykh, 1996. Late Cretaceous
and Paleocene vertebrate paleontology and stratigraphy in the North-eastern
Aral Sea region, Kazakhstan. Journal of Vertebrate Paleontology. 16(3), 46A.
undescribed Tyrannosauroidea (Nessov, 1995)
Early Turonian, Late Cretaceous
Beshtyubin Formation, Kazakhstan
Comments- Nessov (1995) referred material to cf. Alectrosaurus sp.,
though this is doubtful as the numerous other supposed occurences of this genus
in Kazakhstan and Uzbekistan are incorrect.
Reference- Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o
sostave kompleksov, ekologii i paleobiogeografii. Institute for Scientific Research
on the Earth's Crust, St. Petersburg State University, St. Petersburg. 156 pp.
unnamed Tyrannosauroidea (Nessov, 1995)
Turonian-Coniacian, Late Cretaceous
Zhirkindek Formation, Kazakhstan
Material- (ZIN PH 5/49) posterior dorsal neural spine (Averianov, 2007)
(ZIN PH 15/49) tooth fragment (Averianov, 2007)
teeth (Nessov, 1995)
Comments- Nessov (1995) referred teeth from this formation to Alectrosaurus(?),
though this taxon is not known from teeth. Averianov noted the presence of a
tooth fragment and described and illustrated a neural spine.
References- Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o
sostave kompleksov, ekologii i paleobiogeografii. Institute for Scientific Research
on the Earth's Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern
Aral Sea region, Kazakhstan. Cretaceous Research. 28, 532-544.
unnamed Tyrannosauroidea (Shilin and Romanova, 1978)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- (N 485/12457) tooth (Nessov, 1995)
(ZIN PH 10/49) tooth (>80 mm) (Averianov, 2007)
(ZIN PH 11-14/49) tooth fragments (Averianov, 2007)
(ZIN PH 16/49) tooth (Averianov, 2007)
(ZIN PH 17/49) tooth fragment (Averianov, 2007)
(ZIN PH 18/49) tooth fragment (Averianov, 2007)
(ZIN PH 19-22/49) tooth fragments (Averianov, 2007)
teeth (Dyke and Malakhov, 2004)
Comments- Tyrannosauroid material including teeth was first reported
by Shilin and Romanova (1978), and followed by Nessov (1995) and Kordikova et
al. (1996). Dyke and Malakhov (2004) and Averianov (2007) both referred and
illustrated teeth and tooth fragments, though only the latter described them.
Both Kordikova et al. and Dyke and Malakhov referred the material to cf. Alectrosaurus
sp., but teeth are unknown for that genus. The teeth described by Averianov
are distinctive in having a high DSDI (1.31). A femur (N 601/12457) referred
to Tarbosaurus by Nessov (1995) was reidentified as Neimongosaurus
sp. indet. by Averianov (2007).
References- Shilin and Romanova, 1978. [Senonian floras of Kazakhstan].
Alma-Ata, Nauka. 176 pp.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii. Institute for Scientific Research on the Earth's
Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Kordikova, Gunnell, Polly and Kovrizhnykh, 1996. Late Cretaceous and Paleocene
vertebrate paleontology and stratigraphy in the northeastern Aral Sea region,
Kazakhstan. Journal of Vertebrate Paleontology. 16(3), 46A.
Dyke and Malakhov, 2004. Abundance and taphonomy of dinosaur teeth and other
vertebrate remains from the Bostobynskaya Formation, north-east Aral Sea region,
Republic of Kazakhstan. Cretaceous Research. 25(5), 669-674.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern
Aral Sea region, Kazakhstan. Cretaceous Research. 28, 532-544.
undescribed Tyrannosauroidea (Efremov, 1944)
Santonian-Early Campanian, Late Cretaceous
Kara-Cheku, Almaty, Kazakhstan
Comments- This material may belong to the derived tyrannosaurine represented
by dentary IZK 33/MP-61.
Reference- Efremov, 1944. [Dinosaur horizon of Middle Asia and some questions
of stratigraphy]. Izvestiya Akademii Nauk SSSR, Seriya Geologicheskaya. 3, 40-58.
undescribed Tyrannosauroidea (Rozhdestvensky, 1977)
Early Santonian, Late Cretaceous
Yalovach Formation, Tajikistan
Material- teeth
Comments- Nessov (1995) referred to three taxa (10-11 m Carnosauria,
cf. Alectrosaurus sp. (with laterally flattened teeth) and Tyrannosauridae
with relatively thick teeth), all of which are probably tyrannosauroids and
may represent ontogenetic and/or positional variation instead of taxonomic variation.
None are likely to be Alectrosaurus, which isn't known from teeth.
References- Rozhdestvensky, 1977. [Kansai locality of Cretaceous vertebrates
in Fergana]. Yezhyegodnik Vsyesoyuznogo palyeontologichyeskogo obshchyestva.
20, 235-247.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii. Institute for Scientific Research on the Earth's
Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Alifanov and Averianov, 2006. On the finding of ornithomimid dinosaurs (Saurischia,
Ornithomimosauria) in the Upper Cretaceous beds of Tajikistan. Paleontological
Journal. 40(1), 103-108.
undescribed Tyrannosauroidea (Nessov, 1995)
Early Cenomanian, Late Cretaceous
Khodzakul Formation, Uzbekistan
Material- nine teeth
Comments- Nessov (1995) mentions relatively small flattened teeth he
calls Laelaps cf. explanatus, and states may be a peculiar species of
Alectrosaurus. The former is a dromaeosaurid however, and the latter
is not known from teeth. Averianov and Sues (2012) mention Khodzakul tyrannosauroid
teeth which do not differ from Bissekty teeth.
Reference- Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o
sostave kompleksov, ekologii i paleobiogeografii. Institute for Scientific Research
on the Earth's Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Averianov and Sues, 2012. Skeletal remains of Tyrannosauroidea (Dinosauria:
Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan.
Cretaceous Research. 34, 284-297.
undescribed possible Tyrannosauroidea (Leshchinskiy, Voronkevich, Fayngertz,
Maschenko, Lopatin and Averianov, 2001)
Albian?, Early Cretaceous
Shestakovo, Russia
Reference- Leshchinskiy, Voronkevich, Fayngertz, Maschenko, Lopatin and
Averianov, 2001. Early Cretaceous vertebrate locality Shestakovo, Western Siberia,
Russia: A refugium for Jurassic relicts? Journal of Vertebrate Paleontology.
21(3), 73A.
unnamed tyrannosauroid (Perle, 1977)
Cenomanian-Turonian, Late Cretaceous
Bayanshiree Formation, Mongolia
Material- (IGM 100/50) partial maxilla, nasal, three dorsal vertebrae,
seventeen caudal vertebrae, scapulocoracoid, proximal humerus, manual ungual
I
(IGM 100/51) premaxilla, partial maxilla, postorbital, jugal, quadratojugal,
dentary, partial ilium, femur, incomplete tibia, metatarsus
Diagnosis- (after Carr, 2005) metatarsal III pinched out for half its
length posterior to metatarsals II and IV; (after Currie, 2001) first two or
three maxillary teeth incisiform.
Comments-
This was first mentioned by Barsbold (976) as "new materials from the
MPR*", "bones of the recently found alectrosaur from Bayshin Tsav
(Bainsheire Formation, Cenomanian-Lower Cenon) there isa small
ungual phalanx of the first manual digit, quite typical for
tyrannosaurids." Perle (1977) referred this material to Alectrosaurus olseni,
which has been generally followed in the literature. Holtz (2001) found that
it was the sister taxon to A. olseni in an unpublished cladistic analysis.
Carr (2005) found it differs from Alectrosaurus in a few features- hypertrophied
manual flexor tubercles, the entire distal end of metatarsal III is widened
relative to the rest of the bone, and metatarsal III is apomorphically pinched
out for half its length posterior to metatarsals II and IV. He finds no reason
to refer the specimens to Alectrosaurus. Restudy of the material is clearly
necessary.
References-
Barsbold, 1976. New data on Therizinosaurus (Therizinosauridae, Theropoda). In Kramarenko, Luvsandansan, Voronin, Barsbold, Rozhdestvensky, Trofimov and Reshetov (Eds.).
Paleontology and Biostratigraphy of Mongolia. The Joint Soviet-Mongolian
Paleontological Expedition, Transactions. 3, 76-92.
Perle, 1977. On the first discovery of Alectrosaurus
(Tyrannosauridae, Theropoda) from the Late Cretaceous of Mongolia. Problemy
Geologii Mongolii. 3, 104-113.
Currie, 2001. Theropod dinosaurs from the Cretaceous of Mongolia. In Benton,
Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia.
434-455.
Holtz, 2001. Pedigree of the tyrant kings: New information on the origin and
evolution of the Tyrannosauridae. Journal of Vertebrate Paleontology. 21(3),
62A-63A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
undescribed Tyrannosauroidea (Barsbold, Kobayashi and Kubota, 2007)
Cenomanian-Turonian, Late Cretaceous
Bayanshiree Formation, Mongolia
Reference- Barsbold, Kobayashi and Kubota, 2007. New discovery of dinosaur
fossils from the Upper Cretaceous Bayanshiree Formation of Mongolia. Journal
of Vertebrate Paleontology. 27(3), 44A.
undescribed tyrannosauroid (Ford and Chure, 2001)
Cenomanian-Campanian, Late Cretaceous
Bayanshiree or Baruungoyot Formations, Mongolia
Material- (PENN AN SSR) teeth, fragmentary skeleton
Reference- Ford and Chure, 2001. Ghost lineages and the paleogeographic
and temporal distribution of tyrannosaurids. Journal of Vertebrate Paleontology.
21(3), 50A-51A.
undescribed Tyrannosauroidea (Ford and Chure, 2001)
Campanian, Late Cretaceous
Baruungoyot Formation, Mongolia
Material- (PEN AN SSR coll.) teeth, fragmentary skeleton
(ZPAL coll.) teeth, fragmentary skeleton
Reference- Ford and Chure, 2001. Ghost lineages and the paleogeographic
and temporal distribution of tyrannosaurids. Journal of Vertebrate Paleontology.
21(3), 50A-51A.
undescribed tyrannosauroid (Watabe and Tsogtbaatar, 2004)
Late Campanian, Late Cretaceous
Khongil, Djadochta Formation, Mongolia
Material- (000812 Kh-West No.1) femur
....(000812 Kh-West No.2) metatarsal
....(000812 Kh-West No.3) metatarsal
....(000812 Kh-West No.4) vertebra
....(000812 Kh-West No.5) rib
....(000812 Kh-West No.1) supraorbital
Comments- Watabe and
Tsogtbaatar (2004) say "the limb and cranial bones of a large carnosaur
were found" at "the western area of Khovil" on July 30 2000.
However, this is a typo for Khongil, as Yaagan Khovil was explored on
July 21, and the authors later state on August 12 "the excavation work
of large carnosaur isolated bone elements discovered onj July 30 was
carried out in the western are of Khngil. Two metatarsals, a femur, a
vertebra, 1 rib and a supraorbital were collected." The
supraorbital confirms this is a tyrannosaurid.
Reference-
Watabe and Tsogtbaatar, 2004. Report on the Japan - Mongolia Joint Paleontological Expedition to the
Gobi desert, 2000. Hayashibara Museum of Natural Sciences Research
Bulletin. 2, 45-67.
undescribed possible tyrannosauroid (Watabe and Tsogtbaatar, 2004)
Early Maastrichtian, Late Cretaceous
Yagaan Khovil,
Nemegt Formation, Mongolia
Material- (uncollected?) caudal vertebra
(uncollected?) phalanges
Comments- Watabe and
Tsogtbaatar (2004) report "a caudal vertebra of a large carnosaur"
found on July 21 2000 at Yagaan Khovil. While it is likely this
is tyrannosaurid, and thus probably Tarbosaurus, it may be Deinocheirus or Therizinosaurus as well. They also state "large carnosaur digits" were discovered.
Reference-
Watabe and Tsogtbaatar, 2004. Report on the Japan - Mongolia Joint Paleontological Expedition to the
Gobi desert, 2000. Hayashibara Museum of Natural Sciences Research
Bulletin. 2, 45-67.
unnamed possible tyrannosauroid (Young, 1942)
Tithonian?, Late Jurassic
IVPP locality 47, upper Guangyuan Group, Sichuan, China
Material-
(IVPP V237C; syntype of Chienkosaurus ceratosauroides) premaxillary tooth (~13x10x7 mm)
Comments- The material was discovered in late Spring 1941, part of the Chienkosaurus ceratosauroides 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. Dong et al. (1983) 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.
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, IVPP
V237 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.)".
References- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan,
China. Bulletin of the Geological Society of China. 22(3-4), 293-309.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.
undescribed possible tyrannosauroid (Xu, Zheng and Yu, 2010)
Early Cretaceous?
Liaoning, China
Material- (STM coll.) (large) specimen including skull, caudal vertebrae,
feathers
Comments- Xu et al. (2010) mention a large possible tyrannosauroid. It
has broad non-branched feathers ~10 mm wide attached to its caudal vertebrae.
In a newspaper article on Xu's work, Branigan (2011) noted it had "huge,
shark-like teeth and a lengthy tail."
References- Xu, Zheng and Yu, 2010. Exceptional dinosaur fossils show
ontogenetic development of early feathers. Nature. 464, 1338-1341.
Branigan, 2011. Chinese 'dinosaur city' reshapes understanding of prehistoric
era. Guardian. May 14th, 23.
undescribed possible Tyrannosaurioidea (Gilmore, 1933)
Cenomanian-Santonian, Late Cretaceous
‘Nantienmen’ beds, Hebei, China
Material- (AMNH 2906) (~3 m) partial dorsal vertebrae, incomplete sacrum,
partial caudal vertebrae, proximal radius, fragmentary ilia, fragmentary pubes,
fragmentary ischia (Gilmore 1933)
?...(AMNH 6592) (~3 m) tooth, partial cervical vertebra, distal ungual, proximal
ulna, phalanges, partial pubes (Gilmore 1933)
Comments- These specimens derive from the same locality and horizon and
may be from the same individual. The tooth is serrated distally, suggesting
a tyrannosauroid or dromaeosaurid when the age is taken into account. The remains
are illustrated on the AMNH website, where they are identified as tyrannosaurid.
Perhaps it is a juvenile.
Reference- Gilmore, 1933. Two new dinosaurian reptiles from Mongolia
with notes on some fragmentary specimens. American Museum Novitates 679 1-20.
undescribed Tyrannosauridae (Hone, Wang, Sullivan, Zhao, Chen, Li, Ji,
Ji and Xu, 2011)
Campanian, Late Cretaceous
Upper Xingezhuang Formation of the Wangshi Series, Shandong, China
Material- (NGMC V287) tooth fragment
(ZCDM coll.) postcrania
Comments- This may belong to Zhuchengotyrannus or the undescribed
tyrannosaurid (ZCDM V0030 and V0032), but are not described yet. Note the indeterminate
Tyrannosaurus? zhuchengensis is also from the same deposits.
Reference- Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011.
A new, large tyrannosaurine theropod from the Upper Cretaceous of China. Cretaceous
Research. 32(4), 495-503.
undescribed Tyrannosauridae (Gilmore, 1933)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (AMNH 21552) (large) femur (Mader and Bradley, 1989)
(AMNH coll.) (large) pedal elements (Gilmore 1933)
(IVPP coll.) teeth and/or elements (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)
Comments- Under Deinodontidae, Gilmore (1933) states "The presence of a second carnivore, apparently rivaling Tyrannosaurus
in size, is indicated by a few scattered foot bones." These are no
doubt one or more of the specimens listed here under undescribed
Averostra (e.g. AMNH 6376, 6556, 6744, 6756, 6757, etc.).
Mader and Bradley (1989) noted "among the materials brought back by the
Central Asiatic Expeditions was the isolated femur (AMNH 21552) of a
much larger tyrannosaur [than the Alectrosaurus lectotype]." This may belong to the same taxon as Gilmore's pedal material based on size.
Yao et al. (2015) note "small unarticulated bones and teeth, including
fossils of ... tyrannosauroids" from "a rare microvertebrate locality
within the Iren Dabasu Formation, about 16 km northeast of Erenhot
City."
References- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.
Mader and Bradley, 1989. A redescription and revised diagnosis of the syntypes
of the Mongolian tyrannosaur Alectrosaurus olseni. Journal of Vertebrate
Paleontology. 9(1), 1-55.
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. Caenagnathasia
sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous:
Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.
undescribed tyrannosaurid (Russell, Russell and Sweet, 1993)
Late Maastrichtian, Late Cretaceous
Pingling Formation of the Nanxiong Group, Guangdong, China
Reference- Russell, Russell and Sweet, 1993. The end of the dinosaurian
era in the Nanxiong Basin. Vertebrata PalAsiatica. 31(2), 139-145.
unnamed possible tyrannosauroid (Manabe 1999)
Aptian, Early Cretaceous
Jobu Formation of the Itoshiro Subgroup of the Tetori Group, Japan
Material- (IBEF VP 001) premaxillary tooth (11x4.5x3.8 mm)
Description- serrated, D-shaped cross section, posterior surface flat
without central ridge, twenty serrations per 5 mm on both carinae, serrations
comparatively larger than in Gorgosaurus teeth.
Habitat- The specimen was transported from a river side to a lake based
on the condition of the bones. Other inhabitants of the lake included crocodiles,
turtles and fish.
References- Azuma, 1991. Early Cretaceous dinosaur Fauna from the Tetori
Group, central Japan. Research on Dinosaurs from the Tetori Group (1). Professor
S. Miura Memorial Volume, 55-69
Manabe, 1999. The early evolution of the Tyrannosauridae in Asia. Journal of
Paleontology. 73(6), 1176-1178.
undescribed Tyrannosauridae (Jiji Press, 2015)
Campanian, Late Cretaceous
Nagasaki, Kyushu, Japan
Material- (Nagasaki Science Museum coll.) tooth (72x38x27 mm)
(Nagasaki Science Museum coll.) incomplete tooth (?x37x22 mm)
Reference- Jiji Press, 2015. Large tyrannosaurids existed in Japan. The
Japan News. 7-15-2015. http://the-japan-news.com/news/article/0002287344 [not archived]
undescribed possible tyrannosauroid (Naish, DML 2000)
Late Kimmeridgian-Tithonian, Late Jurassic
Tendaguru Formation, Tanzania
Material- (NHMUK coll.) premaxillary tooth (~10 mm)
Description- D-shaped; one side serrated, the other not.
Reference- Naish, DML 2000. https://web.archive.org/web/20191009075255/http://dml.cmnh.org/2000Apr/msg00440.html
unnamed possible tyrannosauroid (Benson, Barrett, Rich and Vickers-Rich,
2010)
Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P186046) pubes (307 mm)
Comments- This is probably the pubis mentioned by Currie et al. (1996)
as NMV P186058, which they referred to Ornithomimosauria. It was later described
by Benson et al. (2010) as a tyrannosauroid closer to tyrannosaurids than Guanlong,
Juratyrant or Raptorex. Herne et al. (2010) argued it could only
be identified as Avetheropoda indet., while Novas et al. (2013) suggested a
megaraptoran affinity.
References- Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur
(Theropoda, Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur
Cove, Australia. Alcheringa. 20(1-2), 73-79.
Benson, Barrett, Rich and Vickers-Rich, 2010. A Southern tyrant reptile. Science.
327(5973), 1613.
Benson, Barrett, Rich, Vickers-Rich, Pickering and Holland, 2010. Response to
Comment on "A Southern tyrant reptile". Science. 329(5995), 1013d.
Herne, Nair and Salisbury, 2010. Comment on "A Southern tyrant reptile".
Science. 329(5995), 1013c.
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.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Bistahieversor Carr and Williamson,
2010
= "Bistahieversor" Carr, 2005
B. sealeyi Carr and Williamson, 2010
= "Bistahieversor sealeyi" Carr, 2005
Late Campanian, Late Cretaceous
Hunter Wash Member of the Kirtland Formation, New Mexico, US
Holotype- (NMMNH P-27469) (adult) skull (1.07 m), mandibles, incomplete
postcranial skeleton including vertebrae, ribs, pelvis, hindlimbs
Late Campanian, Late Cretaceous
Farmington Member of the Kirtland Formation, New Mexico, US
Paratype- (NMMNH P-25049) (1.7 m high at hips, 278 kg, juvenile) incomplete
skull (premaxillary fragment, maxilla, nasals, partial lacrimals, partial jugal,
frontals, parietals, partial postorbital, quadratojugal, quadrate, palatine,
partial ectopterygoids, pterygoid fragment?, parasphenoid, basisphenoid, basioccipital,
laterosphenoid, prootic, exoccipital-opisthotic), partial dentary, surangular
fragment, articular, stapes, partial hyoid, sixteen caudal vertebrae, eight
chevrons, scapula, partial forelimb, partial ilium, femur, tibia, fibula, astragalus,
metatarsus, pes
Late Campanian, Late Cretaceous
Fruitland Formation, New Mexico, US
Paratype- (NMMNH P-32824) partial lacrimal
Late Campanian, Late Cretaceous
Upper Fruitland or Lower Kirtland Formation, New Mexico, US
Paratype- (OMNH 10131) (juvenile) premaxillary tooth (52 mm), maxillary
tooth (75 mm), partial frontal, partial parietal, incomplete postorbital, partial
dentary, four rib fragments, gastralium, distal half of pubis, femur lacking
distal end (~1.033 m), distal half of tibia (~891 mm), distal half of metatarsal
III (~ 483 mm), metatarsal IV (461 mm)
Diagnosis- (after Carr and Williamson, 2010) forked palatal process of
premaxilla; supernumerary frontal processes of nasal; lanceolate medial frontal
processes of nasal; pneumatic foramen that pierces the supraorbital ramus of
lacrimal; peaked sagittal crest; supratemporal fossa extends onto lateral surface
of squamosal; short prefrontal; single pneumatic foramen in palatine; medial
ridge on angular for insertion into the surangular; ventrolateral keel along
posteroventral margin of the mandible formed by angular and prearticular; tall
flange extending from ventral margin of anterior mylohyoid foramen of splenial.
Comments- Carr first named and described this taxon in his unpublished
thesis (Carr, 2005). Lehman and Carpenter previously identified OMNH 10131 as
Aublysodon cf. mirandus, and it was later identified as Daspletosaurus
sp. by Carr and Williamson (2000). Carr and Williamson (2000) previously
identified NMMNH P-25049 as a new species of Daspletosaurus. Carr and
Williamson (2002) and Carr (2005) found Bistahieversor to be the sister
taxon of Tyrannosauridae based on cranial characters. Carr and Williamson later
(2010) officially described the taxon and using a matrix similar to Carr's (2005)
but with more postcranial characters, found it to be in a polytomy with Dryptosaurus,
Appalachiosaurus, Alioramus and Tyrannosauridae. More recently,
Brusatte et al. (2010) found it to be sister to Appalachiosaurus+Tyrannosauridae.
References- Lehman and Carpenter, 1990. A partial skeleton of the tyrannosaurid
dinosaur Aublysodon from the Upper Cretaceous of New Mexico. Journal
of Paleontology. 64, 1026-1032.
Carr and Williamson, 1999. A new tyrannosaurid (Theropoda: Coelurosauria) from
the San Juan Basin of New Mexico. Journal of Vertebrate Paleontology. 19(3),
36A.
Williamson and Carr, 1999. A new tyrannosaurid (Dinosauria: Theropoda) partial
skeleton from the Upper Cretaceous Kirtland Formation, San Juan Basin, New Mexico.
New Mexico Geology, Guidebook 43. 26-29.
Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria: Coelurosauria)
from New Mexico. In Lucas and Heckert (eds.). New Mexico Museum of Natural History
and Science Bulletin. 17, 113-146.
Williamson and Carr, 2001. Dispersal of pachycephalosaurs and tyrannosauroids
between Asia and North America. Journal of Vertebrate Paleontology. 21(3), 114A.
Carr and Williamson, 2002. Evolution of basal Tyrannosauroidea of North America.
Journal of Vertebrate Paleontology. 22(3), 41A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Carr and Williamson, 2010. Bistahieversor sealeyi, gen. et sp. nov.,
a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea.
Journal of Vertebrate Paleontology. 30(1), 1-16.
Magana, D'amore, Molnar and Hall, 2013. Identifying isolated shed teeth from
the Kirtland Formation of northwestern New Mexico. Journal of Vertebrate Paleontology.
Program and Abstracts 2013, 169.
Brusatte, Muir, Averianov, Balanoff, Bever, Carr, Kundrat, Sues,
Williamson and Xu, 2016. Brains before brawn: Neurosensory evolution in
tyrannosauroid dinosaurs. Journal of Vertebrate Paleontology. Program
and Abstracts, 106.
McKeown, Brusatte, Williamson, Schwab, Carr, Butler, Muir, Schroeder,
Espy, Hunter, Losko, Nelson, Gautier and Vogel, 2020. Neurosensory and
sinus evolution as tyrannosauroid dinosaurs developed giant size:
Insight from the endocranial anatomy of Bistahieversor sealeyi. The Anatomical Record. 303(4), 1043-1059.
Carr and Williamson, in prep. Phylogeny of the Tyrannosauroidea.
Appalachiosaurus
Carr, Williamson and Schwimmer, 2005
= "Appalachiosaurus" Holtz, Molnar and Currie, 2004
A. montgomeriensis Carr, Williamson and Schwimmer, 2005
Middle Campanian, Late Cretaceous
Demopolis Formation, Alabama, US
Holotype- (RMM 6670) (623 kg) premaxillary tooth, maxilla, nasals, lacrimal,
partial jugal, palatine, ectopterygoid, incomplete pterygoid, dentary, splenial,
angular, nine lateral teeth (24, 30.4 mm), four proximal caudal vertebrae (94,
88/105 mm), proximal caudal neural arch, incomplete distal caudal vertebra (106
mm), partial distal caudal centrum, pubic shaft, ischium (>496 mm), femora
(786, 754.7 mm), tibiae (763.5, ~780.7 mm), fibulae (~678 mm), astragali (155.1
mm wide), calcanea, metatarsal II (455.8, 458.7 mm), phalanx II-1 (131.5 mm),
phalanx II-2 (94.5 mm), metatarsal III (~482.2 mm), phalanx III-1 (124.9 mm),
phalanx III-2 (92.6 mm), pedal ungual III, metatarsal IV (468.7 mm), phalanx
IV-1 (92.1 mm), phalanx IV-2 (77.4, 78.7 mm), phalanx IV-1 (41.1 mm), pedal
ungual IV
Middle Campanian, Late Cretaceous
Coachman Formation, South Carolina, US
?(ChM PV7326) incomplete tooth (Schwimmer, Sanders, Erickson and Weems, 2015)
?(ChM PV7370) limb fragment (Schwimmer, Sanders, Erickson and Weems, 2015)
?(ChM PV8826) tooth (Schwimmer, Sanders, Erickson and Weems, 2015)
?(ChM PV9117) incomplete tooth (Schwimmer, Sanders, Erickson and Weems, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Donaho Creek or Peedee Formation, South Carolina, US
Material- ?(SCSM 98.64.2) tooth (Schwimmer, Sanders, Erickson and Weems,
2015)
Late Maastrichtian, Late Cretaceous
Steel Creek Formation equivalent, South Carolina, US
Material- ?(ChM PV6819) pedal phalanx (Schwimmer, Sanders, Erickson and
Weems, 2015)
Diagnosis- (after Carr et al., 2005) wide jugal process of ectopterygoid;
caudal pneumatic recess of palatine situated rostral to caudal margin of vomeropterygoid
process; articular surface for lacrimal of palatine situated distally; and prominent
lip extending over dorsal margin of articular surface of pedal unguals.
Comments- The name "Appalachiosaurus" was first used online
by Holtz et al. (2004) in the data matrix of their phylogenetic analysis. None
of the South Carolinean material assigned to Appalachiosaurus by Schwimmer
et al. (2015) was done so based on anatomy, and the later ages of SCSM 98.64.2
and ChM PV6819 suggest they belong to other taxa.
Holtz (2004) found this taxon to be a basal albertosaurine, but after adding
Dilong to his matrix (2005 Burpee Symposium), Appalachiosaurus
ended up basal to Tyrannosauridae, as in Carr et al.'s (2005) and Carr's (2005)
analyses using cranial characters. Most recently, Brusatte et al. (2010) found
it to be sister to Tyrannosauridae.
References- Schwimmer and Kiernan, 2001. Eastern Late Cretaceous theropods
in North America and the crossing of the Interior Seaway. Journal of Vertebrate
Paleontology. 21(3) 99A.
Williamson and Carr, 2001. Dispersal of pachycephalosaurs and tyrannosauroids
between Asia and North America. Journal of Vertebrate Paleontology. 21(3) 114A.
Carr and Williamson, 2002. Evolution of basal Tyrannosauroidea from North America.
Journal of Vertebrate Paleontology. 22(3) 41A.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Carr, Williamson, and Schwimmer, 2005. A new genus and species of tyrannosauroid
from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama.
Journal of Vertebrate Paleontology. 25(1), 119-143.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Schwimmer, Sanders, Erickson and Weems, 2015. A Late Cretaceous dinosaur and
reptile assemblage from South Carolina, USA. Transactions of the American Philosophical
Society. 105(2), 157 pp.
Carr and Williamson, in prep. Phylogeny of the Tyrannosauroidea.
Tyrannosauridae Osborn, 1906
Definition- (Gorgosaurus libratus + Albertosaurus sarcophagus
+ Daspletosaurus torosus + Tarbosaurus bataar + Tyrannosaurus
rex) (Holtz, 2004)
Other definitions- (Tyrannosaurus rex <- Alectrosaurus olseni,
Aublysodon mirandus, Nanotyrannus lancensis) (modified from Sereno, 1998)
(Aublysodon mirandus + Tyrannosaurus rex) (modified from Holtz,
2001)
(Alectrosaurus olseni + Gorgosaurus libratus + Albertosaurus
sarcophagus + Daspletosaurus torosus + Alioramus remotus +
Tarbosaurus bataar + Tyrannosaurus rex) (Brochu, 2003)
(Tyrannosaurus rex <- Eotyrannus lengi) (Holtz, 2004)
(Gorgosaurus libratus + Albertosaurus sarcophagus + Tyrannosaurus
rex) (Sereno et al., 2009)
(Gorgosaurus libratus + Tyrannosaurus rex) (Sereno et al., 2005
vide Brusatte et al., 2011)
= Deinodontidae Cope, 1866 emmend. Brown, 1914
= Aublysodontidae Nopsca, 1928
= Shanshanosauridae Dong, 1977
= Tyrannosauridae sensu Sereno et al. 2009
Definition- (Gorgosaurus libratus + Albertosaurus sarcophagus
+ Tyrannosaurus rex)
= Tyrannosauridae sensu Sereno et al., 2005 vide Brusatte et al., 2011
Definition- (Gorgosaurus libratus + Tyrannosaurus rex)
Comments- While unnamed specimens from the Campanian and Maastrichtian
are listed below as tyrannosaurids, the presence of late surviving taxa such
as Alectrosaurus and Dryptosaurus suggests many may be more basal
tyrannosauroids.
At least one family has precedence over Tyrannosauridae- Deinodontidae (originally
misspelled Dinodontidae by Cope) from 1866, which is based on the genus Deinodon.
Deinodon consists of several teeth of dubious association which are probably
referrable to Gorgosaurus and/or Daspletosaurus. Deinodontidae
was commonly used before the 1950's and Tyrannosauridae was mostly used after
1970, perhaps based on Russell (1970). Russell described Deinodon as
a 'nomen vanum' (= nomen dubium) and stated it "is not a useful systematic
procedure to perpetuate family group names based on generically unidentifiable
material", but this is not a rule in the ICZN and the genus has not been
reevaluated recently.
Tyrannosauridae defined- Sereno's (1998) definition of Tyrannosauridae
is problematic, as Nanotyrannus is probably a junior synonym of Tyrannosaurus
and it seems likely Aublysodon is a tyrannosaurine. The latter also means
Holtz's (2001) definition would only include tyrannosaurines. Brochu's (2003)
definition includes Alectrosaurus, which is here resolved as far more
basal within Tyrannosauroidea. Holtz (2004) gave Tyrannosauridae two different
definitions in his Dinosauria chapter, presumably on accident. One is stem-based
and would make the family cover several more basal taxa like Bistahieversor,
Appalachiosaurus, Dryptosaurus and Xiongguanlong (notably
Dryptosauridae has priority over Tyrannosauridae). The other is node-based and
is used here. Sereno et al.'s (2009) definition is a first order redefinition
of Holtz's second definition, after deleting Daspletosaurus and Tarbosaurus.
I agree with Sereno that their inclusion is useless, as none are ever placed
outside (Gorgosaurus + Albertosaurus + Tyrannosaurus),
yet their exclusion is also useless. Brusatte et al. (2011) incorrectly attributed
a definition to Sereno et al. (2005) (perhaps intending the 2009 paper), but
did not use Albertosaurus as an internal specifier.
References- Cope, 1866. [On the remains of a gigantic extinct dinosaur,
from the Cretaceous Green Sand of New Jersey]. Proceedings of the Academy of
Natural Sciences of Philadelphia. 18, 275-279.
Osborn, 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur (Second
communication). Bulletin of the American Museum of Natural History. 22(16),
281-296.
Brown, 1914. Cretaceous Eocene correlations in New Mexico, Wyoming, Montana.
Bulletin of the Geological Society of America. 25, 355-380.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Russell, 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National
Museum of Natural Sciences, Publications in Palaeontology. 1, 1-34.
Dong, 1977. On the dinosaurian remains from Turpan, Xinjiang. Vertebrata PalAsiatica.
15(1), 59-66.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210(1), 41-83.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. In Tanke and
Carpenter (eds.). Mesozoic Vertebrate Life. 64-83.
Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from a nearly
complete skeleton and high-resolution computed tomographic analysis of the skull.
SVP Memior 7. 138 pp.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Snively and Henderson, 2004. Nasal fusion reinforced the rostrum of tyrannosaurids.
Journal of Vertebrate Paleontology. 24(3), 179A.
Shychoski, 2006. Geometric morphometric, eigenshape and finite element analysis
of cranial variation in tyrannosaurid dinosaurs. Journal of Vertebrate Paleontology.
26(3), 125A.
Shychoski and Snively, 2008. Ecological implications of tyrannosaurid lower
jaw ontogeny, biomechanical scaling and bite function. Journal of Vertebrate
Paleontology. 28(3), 142A.
Reichel, 2009. The heterodonty of tyrannosaurids: Biomechanical implications
inferred through 3D models. Journal of Vertebrate Paleontology. 29(3), 170A.
Sereno, Tan, Brusatte, Kriegstein, Zhao and Cloward, 2009. Tyrannosaurid skeletal
design first evolved at small body size. Science. 326(5951), 418-422.
Hwang and Claire, 2010. Species and genus-level variation in the tooth enamel
microstructure of tyrannosaurid dinosaurs. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 109A.
Shychoski, Snively and Burns, 2010. Maneuvered out of a corner: Ligament entheses
of the arctometatarsus enhanced tyrannosaurid agility. Journal of Vertebrate
Paleontology. Program and Abstracts 2010, 165A.
Brusatte, Benson and Norell, 2011. The anatomy of Dryptosaurus aquilunguis
(Dinosauria: Theropoda) and a review of its tyrannosauroid affinities. American
Museum Novitates. 3717, 53 pp.
Carr, 2011. A comparative study of ontogeny between derived tyrannosauroids:
Evidence for heterochrony. Journal of Vertebrate Paleontology. Program and Abstracts
2011, 84.
Hendricks and Erickson, 2014. A biomechanical explanation for the ampullae of
tyrannosaurid teeth based upon fracture mechanics. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 146.
Snively, Russell, Powell, Theodor and Ryan, 2014. The role of the neck in the
feeding behaviour of the Tyrannosauridae: Inference based on kinematics and
muscle function of extant avians. Journal of Zoology. 292(4), 290-303.
Bykowski and Polly, 2016. Ecologiocal ontogeny and functional changes
in the growth of tyrannosauroids. Journal of Vertebrate Paleontology.
Program
and Abstracts,107-108.
"Alamotyrannus" Dalman
and Lucas, in press in Dalman, 2013
"A. brinkmani" Dalman and Lucas, in press in Dalman, 2013
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (AMNH 2359) tooth, tooth fragments (Cope, 1885)
(AMNH 5882) pedal phalanx (Carr and Williamson, 2000)
(NMMNH P-7199) partial dentary, tooth fragments, partial vertebra (Carr and
Williamson, 2000)
(NMMNH P-13000; = UNM FKK-076) tooth (Lucas et al., 1987)
(NMMNH P-28367) tooth fragment (Carr and Williamson, 2000)
(NMMNH P-28368) tooth (Carr and Williamson, 2000)
(NMMNH P-28369) tooth fragment (Carr and Williamson, 2000)
(NMMNH P-30076) tooth fragment (Carr and Williamson, 2000)
(NMMNH P-32566) tooth fragments (Carr and Williamson, 2000)
(NMMNH P-32567) tooth (?x8.7x~5.1 mm) (Carr and Williamson, 2000)
(NMMNH P-32588) partial premaxillary tooth (Carr and Williamson, 2000)
(NMMNH P-32598) partial premaxillary tooth (Carr and Williamson, 2000)
(NMMNH P-32812) incomplete tibia (Williamson and Carr, 2005)
(NMMNH P-32819) tooth (16.9x~8.6x~3.7 mm) (Williamson and Brusatte, 2014)
(NMMNH P-32888) tooth (Williamson and Carr, 2005)
(NMMNH P-33894) partial metatarsal (Willaimson and Carr, 2005)
(Ratkevich coll.) metatarsal IV (Lehman, 1981)
(SMP VP-1113) incomplete femur (~1 m) (Carr and Williamson, 2000)
(SMP VP-1317) tooth fragment (Jasinski, Sullivan and Lucas, 2011)
(SMP VP-1574) tooth (61 mm) (Sullivan, Lucas and Braman, 2005)
(SMP VP-1848) metatarsal I (Jasinski, Sullivan and Lucas, 2011)
(SMP VP-2105) incomplete scapulocoracoid (Jasinski, Sullivan and Lucas, 2011)
(SMP VP-2174) tooth fragment, fragment (Jasinski, Sullivan and Lucas, 2011)
(SMP VP-2352) partial tooth (105 mm) (Jasinski, Sullivan and Lucas, 2011)
(UNM FKK-77) tooth (Lucas et al., 1987)
(UNM FKK-78) tooth (Lucas et al., 1987)
(UNM FKK-79) tooth (Lucas et al., 1987)
(USNM coll.) teeth (Gilmore, 1916)
(USNM coll.) (~12 m) few vertebrae (Gilmore, 1919)
Comments- Gilmore first reported ?Deinodon teeth (Gilmore, 1916),
then noted large theropod vertebrae (Gilmore, 1919). Lehman (1981) reported
a metatarsal and unspecified NMMNH teeth as ?Albertosaurus. Lucas et
al. (1987) referred NMMNH P-13000 to cf. Tyrannosaurus rex and three
UNM teeth to Albertosaurus. Carr and Williamson (2000) referred NMMNH
P-7199 to Tyrannosaurus,and Williamson and Carr (2005) identified NMMNH
P-13000, 32888, 33894, 32812 and AMNH 5882 as cf. Tyrannosaurus rex.
Sullivan et al. (2005) identified the femur SMP V-1113 as Daspletosaurus
sp., based on comparison to a Gorgosaurus or Daspletosaurus sp.
nov. specimen. Jasinski et al. (2011) thought the scapulocoracoid might be Tyrannosaurus
sp., though they listed all the Naashoibito material as Tyrannosauridae indet..
Dalman (2013) stated the Naashoibito material is diagnostic, and provided a
reference to an in press paper naming it "Alamotyrannus brinkmani."
It is unknown which specimen is the intended holotype or what the diagnostic
characters are. Dalman later (pers. comm. to Demirjian, 2015) stated the paper
is postponed as more complete remains were discovered, and that the taxon would
receive a different name.
References- Cope, 1885. The Vertebrata of the Tertiary formations of
the West, Book 1: Reports of the United States Geological Survey Territories,
F. V. Hayden in charge. 3, 1009 pp.
Gilmore, 1916. Vertebrate faunas of the Ojo Alamo, Kirtland and Fruitland formations.
United States Geological Survey, Professional Paper. 98Q, 279-308.
Gilmore, 1919. Reptilian faunas of the Torrejon, Puerco, and underlying Upper
Cretaceous formations of San Juan County, New Mexico. U.S. Geological Survey,
Professional Paper 119, 71 pp.
Lehman, 1981. The Alamo Wash local fauna: A new look at the old Ojo Alamo fauna.
In Lucas, Rigby and Kues (eds.). Advances in San Juan Basin paleontology. University
of New Mexico Press. 189-221.
Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria: Coelurosauria)
from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico
Museum of Natural History and Science. Bulletin. 17, 113-146.
Sullivan, Luvas and Braman, 2005. Dinosaurs, pollen, and the Cretaceous-Tertiary
boundary in the San Juan Basin, New Mexico. New Mexico Geological Society, 56th
Field Conference Guidebook, Geology of
the Chama Basin. 56, 395-407.
Williamson and Carr, 2005. Latest Cretaceous tyrannosaurs from the San Juan
basin, New Mexico. Abstracts of proceedings from "100 years of Tyrannosaurus
rex, a symposium." 38.
Jasinski, Sullivan and Lucas, 2011. Taxonomic composition of the Alamo Wash
local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member),
San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann (eds.). Fossil
Record 3. New Mexico Museum of Natural History and Science Bulletin. 53, 216-271.
Dalman, 2013. New examples of Tyrannosaurus rex from the Lance Formation
of Wyoming, United States. Bulletin of the Peabody Museum of Natural History.
54(2), 241-254.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Dalman and Lucas, in press. A new large tyrannosaurid Alamotyrannus brinkmani,
n. gen., n. sp. (Theropoda: Tyrannosauridae), from the Upper Cretaceous Ojo
Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico
Museum of Natural History and Science Bulletin.
Aublysodon? lateralis Cope,
1876
= Deinodon lateralis (Cope, 1876) Hay, 1902
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Holotype- (AMNH 3956) (adult) anterior dentary tooth (>25 mm)
....(?) anterior tooth (>11 mm)
Comments- Molnar and Carpenter (1989) thought the serrated carinae indicated
this was a lateral premaxillary tooth from Dromaeosaurus, but juvenile
tyrannosaurids can have serrated carinae as well (e.g. CMN 41104) and the larger
tooth is much too large for Dromaeosaurus.
This taxon is based on two teeth, one much larger than the other. Both are clearly
anterior teeth, as the mesial carina is shifted lingually (not laterally as
in Cope's description). The larger one is described in more detail and must
be tyrannosaurid based on size (FABL of 18 mm, compared to 8 mm or less for
other Judith River theropods). The photograph in Glut (1997) resembles dentary
tooth 4 of Tyrannosaurus most closely and is between dentary teeth 4
and 6 of Gorgosaurus in crown compression. It is comparable in size to
adult tyrannosaurids (FABL of Gorgosaurus specimen ROM 1247 is 21 mm;
of Daspletosaurus specimen MOR 590 is 23 mm), so is probably itself from
an adult. The compression (.56) is comparable to Gorgosaurus (.51-.61)
but less than Daspletosaurus (.74-.78), making it probably referrable
to the former taxon. Both carinae are serrated and the photo indicates the apical
portion has been worn away.
The smaller tooth has a FABL of 6 mm, putting it within the size range of Dromaeosaurus
in addition to juvenile tyrannosaurids. It is not described except to note similarity
to the large tooth with the exception of having a less truncated lingual face.
The photo confirms this, but it is merely due to the angle of wear as opposed
to any anatomical difference. It may be another tyrannosaurid anterior dentary
tooth, or perhaps a Dromaeosaurus premaxillary tooth.
References- Cope, 1876. Descriptions of some vertebrate remains from
the Fort Union Beds of Montana. Paleontological Bulletin. 22, 1-14.
Cope, 1876. Descriptions of some vertebrate remains from the Fort Union Beds
of Montana. Proceedings of the Academy of Natural Sciences of Philadelphia.
28, 248-261.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 1-868.
Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian, Montana, U.S.A.)
referred to the genus Aublysodon. Geobios. 22, 445-454.
Glut, 1997. Dinosaurs, the Encyclopedia. Mcfarland & Company, Inc.. 1076
pp.
Deinodontinae sensu Matthew and Brown, 1922
Deinodon Leidy, 1856
D. horridus Leidy, 1856
= Megalosaurus (Deinodon) horridus (Leidy, 1856) Leidy, 1857
= Aublysodon horridus (Leidy, 1856) Cope, 1868
?= Dryptosaurus kenabekides Hay, 1899
?= Deinodon kenabekides (Hay, 1899) Olshevsky, 1995
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Syntypes- (ANSP 9533; paralectotype of Aublysodon mirandus) premaxillary
tooth
?(ANSP 9534; paralectotype of Aublysodon mirandus) first dentary tooth
fragment
Referred- ?(ANSP 9530; syntype of Dryptosaurus kenabekides) partial
lateral tooth (Leidy, 1856)
?(ANSP 9531) first dentary tooth (Leidy, 1856)
?(ANSP 9536; syntype of Dryptosaurus kenabekides) partial lateral tooth
(Leidy, 1856)
?(ANSP 9538) tooth (Leidy, 1856)
?(ANSP 9539) tooth (Leidy, 1856)
?(ANSP 9540) tooth (Leidy, 1856)
?(ANSP 9541; syntype of Dryptosaurus kenabekides) lateral tooth (Leidy,
1856)
?(ANSP 9542; syntype of Dryptosaurus kenabekides) lateral tooth (Leidy,
1856)
?(ANSP 9543; syntype of Dryptosaurus kenabekides) lateral tooth (Leidy,
1856)
?(ANSP 9544) tooth (Leidy, 1856)
Comments- Leidy (1856) based this species on fourteen teeth and tooth
fragments discovered in the Judith River Group of Montana. Most were lateral
teeth he regarded as different from Megalosaurus only in their greater
labiolingual thickness, but Leidy placed species in the new genus Deinodon
because of several other teeth which he felt were distinctive. These were ANSP
9531, 9533, 9534 and 9535, which can all now be recognized as tyrannosaurid
anterior teeth. Leidy later (1857) sunk his own genus into Megalosaurus
as a subgenus to create the short lived combination Megalosaurus (Deinodon)
horridus. Cope (1866) described the teeth of Deinodon as D-shaped,
referencing 9533-9535, to distinguish them from his new taxon Laelaps
(later renamed Dryptosaurus). This makes him first reviser of the genus,
and connected the name Deinodon horridus to the D-shaped teeth in Leidy's
syntype series. Cope considered the lateral teeth to belong to Laelaps.
Leidy (1868) created the new taxon Aublysodon mirandus for ANSP 9533-9535,
intending to retain Deinodon horridus for the lateral teeth (at least
ANSP 9530, 9536 and 9541-9543). Cope's 1866 specification of Deinodon
for the D-shaped teeth has priority though, making Aublysodon mirandus
an objective junior synonym of Deinodon horridus. Later, Cope (1868)
believed Deinodon was preoccupied by the snake genus Dinodon,
and used the name Aublysodon horridus for the anterior teeth (since he
had attached the species name horridus to the teeth in 1866). Yet Hay
(1899) correctly noted the spellings are different, and thus Deinodon
is still valid. Marsh (1892) followed Leidy's (1868) assignment of D-shaped
teeth to Aublysodon, and considered ANSP 9535 to be typical of A.
mirandus, while ANSP 9533 and 9534 were considered examples of another unnamed
Aublysodon species. A. mirandus was notable for its lack of serrations
compared to 9533 and 9534. This made ANSP 9535 the lectotype of Aublysodon,
which was formalized by Carpenter (1982). ANSP 9533 and 9534 are thus implicitly
the remaining syntypes of Deinodon. Hay (1899) realized restricting Deinodon
and/or Aublysodon to the D-shaped teeth meant the lateral teeth were
without a taxon. Based on the resemblence to Dryptosaurus, he made these
teeth the syntypes of Dryptosaurus kenabekides. Matthew and Brown (1922)
synonymized Deinodon horridus with Aublysodon mirandus and Dryptosaurus
kenabekides, and tentatively with Aublysodon lateralis, Laelaps
incrassatus, L. hazenianus, and Ornithomimus grandis. They
viewed Albertosaurus and/or Gorgosaurus as probably being Deinodon
as well. Russell (1970) stated the Deinodon syntypes cannot be distinguished
from Gorgosaurus or Daspletosaurus and the genus is thus a nomen
dubium. As there is no particular taxonomic reason to separate the lateral and
premaxillary teeth (which all may belong to different individuals and taxa in
any case), they are all retained here under Deinodon horridus.
ANSP 9530 (figures 21-24 in Leidy, 1856) consists of two tooth fragments- a
mesial edge, and the tip. It was considered typical of Deinodon horridus
by Leidy (1868), but made a syntype of Dryptosaurus kenabekides by Hay
(1899). Serrations extend from the tip most of the way down the mesial carina
and along the preserved tip of the distal carina. The mesial carina shifts lingually
at its base, as in the second through ninth maxillary teeth and fifth through
eighth dentary teeth of Tyrannosaurus. The estimated crown compression
(~.50) is more similar to Gorgosaurus (.51-.68) than to Daspletosaurus
(>55-.76), so it may belong to Gorgosaurus.
ANSP 9531 (figures 46-48) is a small tooth crown described by Leidy in 1856
and 1860 as different than the majority of Deinodon teeth, but not included
as an Aublysodon syntype in 1868, or necessarily referenced by Cope (1866)
in his revision of Deinodon since it is not D-shaped in the cross section
illustrated. Its taxonomic status is thus a referred specimen of Deinodon
horridus. It is nearly conical, with a compression of .90. Both carinae
are serrated and shifted lingually to form a D-shape apically. The crown itself
is straight in lingual view, though curved slightly lingually. This morphology
compares to the first dentary tooth of tyrannosaurids, but whether it is referrable
to Gorgosaurus or Daspletosaurus is unknown.
ANSP 9533 and 9534 are syntypes of Deinodon horridus, and paralectotypes
of Aublysodon mirandus. ANSP 9533 (figures 37-40) is clearly a tyrannosaurid
premaxillary tooth, being labiolingually wider than mesiodistally long (by 153%)
and D-shaped. Both carinae are serrated and the lingual face is slightly convex.
Lambe (1917) felt it was more robust than Gorgosaurus, but no detailed
comparisons between Gorgosaurus and Daspletosaurus premaxillary
teeth have been made. ANSP 9534 (figures 33-34) consists of a fragment which
has a serrated carina that forms a right angle in section. It may be a second
dentary tooth, as this has a right angled distal carina in Tyrannosaurus.
ANSP 9535 (figures 41-45) is the lectotype of Aublysodon mirandus. As
it lacks serrations, it is a juvenile tyrannosaurine premaxillary tooth and
probably referrable to Daspletosaurus (see Aublysodon entry).
ANSP 9536, 9541, 9542 and 9543 are all syntypes of Dryptosaurus kenabekides
and considered Deinodon horridus by Leidy in 1868.
References- Leidy, 1856. Notices of the remains of extinct reptiles and
fishes, discovered by Dr. F.V. Hayden in the badlands of the Judith River, Nebraska
Territory. Proceedings of the Academy of Natural Sciences of Philadelphia. 8(2),
72-73.
Leidy, 1857. List of extinct Vertebrata, the remans of which have been discovered
in the region of the Missouri River: With remarks on their geological age. Proceedings
of the Academy of Natural Sciences of Philadelphia. 9, 89-91.
Leidy, 1860. Extinct Vertebrata from the Judith River and Great Lignite Formations
of Nebraska. American Philosophical Society Transactions. 11, 139-154.
Cope, 1866. [On the remains of a gigantic extinct dinosaur, from the Cretaceous
Green Sand of New Jersey]. Proceedings of the Academy of Natural Sciences of
Philadelphia. 18, 275-279.
Cope, 1868. On the genus Laelaps. The American Journal of Science, series
2. 46, 415-417.
Leidy, 1868. Remarks on a jaw fragment of Megalosaurus. Proceedings of
the Academy of Natural Sciences of Philadelphia. 1870, 197-200.
Hay, 1899. On the nomenclature of certain American fossil vertebrates. The American
Geologist. 24, 345-349.
Marsh, 1892. Notes on Mesozoic vertebrate fossils. American Journal of Science.
44, 170-176.
Brown, 1914. Cretaceous Eocene correlations in New Mexico, Wyoming, Montana.
Bulletin of the Geological Society of America. 25, 355-380.
Lambe, 1917. The Cretaceous theropodous dinosaur Gorgosaurus. Geological
Survey of Canada, Memoir. 100, 1-84.
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.
Russell, 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National
Museum of Natural Sciences, Publications in Palaeontology. 1, 1-34.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek
formations and a description of a new species of theropod. Contributions to
Geology, University of Wyoming. 20(2), 123-134.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku
Saizensen. 9, 92-119 (part 1); 10, 75-99 (part 2).
Spamer, Daeschler and Daeschler, 1995. A Study of Fossil Vertebrate Types in
the Academy of Natural Sciences of Philadelphia. 434 pp.
Deinodon? falculus (Cope, 1876)
Osborn, 1902
= Laelaps falculus Cope, 1876
= Dryptosaurus falculus (Cope, 1876) Hay, 1902
= Dromaeosaurus falculus (Cope, 1876) Olshevsky, 1978
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Holotype- (AMNH 3959) tooth (9 mm), nine teeth
Comments- The described tooth lacks mesial serrations, as in some examples
of juvenile tyrannosaurids, Richardoestesia and Saurornitholestes.
Crown compression (BW of 4.0 mm / FABL of 5.6 mm) is comparable to tyrannosaurids
and Dromaeosaurus, but outside the range of Saurornitholestes
and Richardoestesia. Crown elongation is similar to all except Richardoestesia.
When compression and elongation are analyzed together, falculus falls
out within Tyrannosauridae, just outside Dromaeosaurus, and far from
Saurornitholestes and Richardoestesia. Similarly, serration size
(at least 5/mm) falls out within tyrannosaurids when plotted against BW, and
within tyrannosaurids and Dromaeosaurus when plotted against crown compression.
When all components are analyzed together, falculus is comparable to
tyrannosaurids and very close to Dromaeosaurus. The evidence suggests
that falculus is a juvenile tyrannosaurid tooth, probably Gorgosaurus
and/or Daspletosaurus based on provenance.
The teeth figured as Laelaps falculus by Glut (1997) are actually AMNH
3968, unnamed tyrannosaurid teeth. The type teeth remain unillustrated.
References- Cope, 1876. Descriptions of some vertebrate remains from
the Fort Union Beds of Montana. Proceedings of the Academy of Natural Sciences
of Philadelphia. 28, 248-261.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 1-868.
Osborn, 1902. On Vertebrata of the Mid-Cretaceous of the Northwest Territory.
I: Distinctive characters of the Mid-Cretaceous fauna. Contributions to Canadian
Palaeontology. 3, 1-21.
Olshevsky, 1978. The archosaurian taxa. Mesozoic Meanderings. 1, 50 pp.
Glut, 1997. Dinosaurs, the Encyclopedia. Mcfarland & Company, Inc.. 1076
pp.
Deinodon? grandis (Marsh, 1890)
Osborn, 1916
= Ornithomimus grandis Marsh, 1890
= Aublysodon grandis (Marsh, 1890) Huene, 1932
Early Campanian, Late Cretaceous
Eagle Sandstone, Montana, US
Holotype- (USNM coll.; lost) (~8 m) metatarsal III (600 mm, 90 mm transversely)
Comments- Discovered in 1888, “fragments representing a considerable
portion of a skeleton” were also reported by Stanton and Hatcher (1905).
Assumed to be a tyrannosauroid based on size, but could be another arctometatarsalian
theropod. Note the two specimens referred to O. grandis by Marsh (1896)
are from the later Lance Formation and are referrable to Tyrannosaurus
(Gilmore, 1920).
References- Marsh, 1890. Description of new dinosaurian reptiles. The
American Journal of Science. 39, 81-86.
Marsh, 1896. The dinosaurs of North America. United States Geological Survey,
16th Annual Report, 1894-95. 55, 133-244.
Stanton and Hatcher, 1905. Geology and paleontology of the Judith River beds.
United States Geological Survey Bulletin. 257, 1-128.
Osborn, 1916. Skeletal adaptations of Ornitholestes, Struthiomimus,
Tyrannosaurus. Bulletin of the American Museum of Natural History. 35(43),
733-771.
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.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1-2), 1-361.
Deinodon? hazenianus (Cope,
1876) Osborn, 1902
= Laelaps hazenianus Cope, 1876
= Dryptosaurus hazenianus (Cope, 1876) Hay, 1902
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Syntypes- (AMNH 3957) (juvenile) posterior tooth (14 mm)
.... (juvenile) six teeth
Comments- Cope (1876) notes the type tooth has a FABL of 11 mm and a
CW of 7 mm. While the thickness limits the teeth to dromaeosaurines and juvenile
tyrannosaurids, the size and shortness is only comparable to the latter. The
serration size is comparable to either, as is the mesial carina twisting lingually(?).
The lack of facets or flattened sides eliminates Zapsalis from consideration.
Sankey et al. (2002) noted that many tyrannosaurid teeth that had twisted mesial
carinae were transitional between premaxillary and maxillary teeth, however
Smith (2005) finds that the shortest and most curved tyrannosaurid crowns are
from the first dentary tooth (which has a different morphology), last two maxillary
teeth and last three dentary teeth. D? hazenianus probably consists of
posterior maxillary and dentary teeth from juvenile tyrannosaurids, probably
Gorgosaurus and/or Daspletosaurus based on the locality. They
are illustrated by Glut (1997).
References- Cope, 1876. On some extinct reptiles and Batrachia from the
Judith River and Fox Hills Beds of Montana. Proceedings of the Academy of Natural
Sciences of Philadelphia. 28, 340-359.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 1-868.
Osborn, 1902. On Vertebrata of the Mid-Cretaceous of the Northwest Territory.
I: Distinctive characters of the Mid-Cretaceous fauna. Contributions to Canadian
Palaeontology. 3, 1-21.
Glut, 1997. Dinosaurs, the Encyclopedia. Mcfarland & Company, Inc.. 1076
pp.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Smith, 2005. Heterodonty in Tyrannosaurus rex: Implications for the taxonomic
and systematic utility of theropod dentitions. Journal of Vertebrate Paleontology.
25(4), 865-887.
"Ornithomimus" tenuis
Marsh, 1890
= Struthiomimus tenuis (Marsh, 1890) Osborn, 1916
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Holotype- (USNM 5814) distal metatarsal III
Comments- Gilmore (1920) illustrated the specimen for the first time
and felt it resembled tyrannosaurids more than ornithomimids. Russell (1972)
considered Ornithomimus tenuis a possible troodontid, though without
comment. However, it is indeed more similar to tyrannosaurids in having an anterior
fossa just proximal to the articular condyle and lacking the proximally extended
articular surface (posteriorly) of troodontids. It's probably a juvenile Gorgosaurus
or Daspletosaurus, based on provenence.
References- Marsh, 1890. Description of new dinosaurian reptiles. The
American Journal of Science. Series 3. 39, 81-86.
Osborn, 1916. Skeletal adaptations of Ornitholestes, Struthiomimus,
Tyrannosaurus. Bulletin of the American Museum of Natural History. 35(43),
733-771.
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.
Russell, 1972. Ostrich dinosaurs of the Late Cretaceous of Western Canada. Canadian
Journal of Earth Sciences. 9, 375-402.
"Suciasaurus" State of Washington 66th Legislature, 2019
"S. rex" State of Washington 66th Legislature, 2019
Middle Campanian, Late Cretaceous
Cedar District Formation of the Nanaimo Group, Washington, US
Material- (UWBM 95770) femoral fragment (~1.2 m)
Comments-
The element was discovered in April 10 2012 and described by Peecook
and Sidor (2015) as Theropoda indet.. The locality, large size and
fourth trochanter indicate it is probably tyrannosaurid, but it is
indeterminate. In 2019 the 66th Legislature of the State of Washington
proposed House Bill 2155 designating a state dinosaur, which stated
"the dinosaur has been nicknamed Suciasaurus rex." This and the
Substitute House Bill 2155 for the 2020 session which made some
adjustments were covered by the media in those years.
References- Peecook and Sidor, 2015. The first dinosaur from Washington
state and a review of Pacific Coast dinosaurs from North America. PLoS ONE.
10(5), e0127792.
House Bill 2155, 66th Legislature, 2019 Regular Session (Washington 2019).
Substitute House Bill 2155, 66th Legislature, 2020 Regular Session (Washington 2020).
Chingkankousaurus Young,
1958
C. fragilis Young, 1958
Campanian-Middle Maastrichtian, Late Cretaceous
Wangshi Series, China
Holotype- (IVPP V636) partial scapula
Diagnosis- (after Brusatte et al., 2013) indeterminate within derived
Tyrannosauroidea.
Comments- This specimen was originally identified as a theropod scapula,
and later assigned to the Tyrannosauridae by Molnar et al. (1990) because of
its narrow shaft. Chure (2000) was uncertain if the element was a scapula based
on the the medial ridge giving it a symmetrical section proximally, but Brusatte
et al. (2013) showed this is normal for theropod scapulae and that at midshaft
the blade has the usual theropod shape of a teardrop wider dorsally. The latter
authors redescribed the specimen and noted numerous similarities with tyrannosaurids.
They assigned it to Tyrannosauroidea more derived than Dilong based on
the narrow blade and highly expanded distal end.
References- Young, 1958. The dinosaurian remains of Laiyang, Shantung.
Palaeontologia Sinica, New Series C. 42(16), 1-138.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 169-209.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis, Columbia University, 964 pp.
Brusatte, Hone and Xu, 2013. Phylogenetic revision of Chingkankousaurus fragilis,
a forgotten tyrannosauroid from the Late Cretaceous of China. In Parrish, Molnar,
Currie and Koppelhus (eds.). Tyrannosaurid Paleobiology. Indiana University
Press. 1-13.
undescribed Tyrannosauridae (Nelms, 1989)
Coniacian-Maastrichtian, Late Cretaceous
Prince Creek Formation, Alaska, US
Material- (UAM-AK83.V90) (Erickson, 1995)
(UAM-AK298.V031) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK300.V086) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK383.V172) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK383.V175) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK390.V034) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK390.V091) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK455.V001) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK461.V001) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK491.V089) tooth (Fiorillo and Gangloff, 2000)
vertebrae (Gangloff, 1998)
tooth tip (Clos, 2004)
cranial, axial and appendicular material (Fiorillo and Tykoski, 2013)
Comments- Nelms (1989) first reported small tyrannosaurid teeth from
the Prince Creek Formation. Most of the teeth in Fiorillo and Gangloff (2000)
are from the Campanian-Maastrichtian Kogosukruk Tongue portion of the Prince
Creek Formation, five of which are Early Maastrichtian in age. Clos (2004) referred
a partial tooth from the Early Maastrichtian to Albertosaurus. This is
possible given its age. Fiorillo and Tykoski's (2013) material is from 3-4 sites
from the Maastrichtian. The Late Maastrichtian material may belong to Nanuqsaurus.
References- Nelms, 1989. Late Cretaceous dinosaurs from the North Slope
of Alaska. Journal of Vertebrate Paleontology. 9 (3), 34A.
Erickson, 1995. Split carinae on tyrannosaurid teeth and implications of their
development. Journal of Vertebrate Paleontology. 15(2), 268-274.
Gangloff, 1998. Arctic dinosaurs with emphasis on the Cretaceous record of Alaska
and the Eurasian-North American connection. In Lucas, Kirkland and Estep (eds.).
Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural
History and Science Bulletin. 14, 211-220.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation
(Cretaceous) of northern Alaska, with speculations on Arctic dinosaur paleoecology.
Journal of Vertebrate Paleontology. 20(4), 675-682.
Clos, 2004. The Cretaceous in North America. Fossil News, Journal of Amateur
Paleontology. 10(11), 4-11, 14-18.
Fiorillo and Tykoski, 2013. Distribution and polar paleoenvironments of large
theropod skeletal remains from the Prince Creek Formation (Early-Late Maastrichtian)
of northern Alaska. Journal of Vertebrate Paleontology. Program and Abstracts
2013, 127.
undescribed Tyrannosauridae (Buckley, McCrea and Currie, 2005)
Late Campanian-Early Maastrichtian, Late Cretaceous
Wapiti Formation, British Columbia, Canada
Material- (PRPRC coll.) partial anterior tooth (Buckley, McCrea and Currie, 2005)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
(UALVP 48760) tooth (Fanti and Miyashita, 2009)
(UALVP 48773.2007.5) ?dentary tooth (?x34.5x30 mm) (Fanti and Miyashita, 2009)
(UALVP 50641.01) tooth (Fanti and Miyashita, 2009)
(UALVP 50641.02) premaxillary tooth (Fanti and Miyashita, 2009)
(UALVP 52580) premaxillary tooth (Fanti, Currie and Burns, 2015)
forty-nine teeth (Fanti and Miyashita, 2009)
eleven teeth (Fanti, Currie and Burns, 2015)
Comments- Buckley et al. (2005) referred the British Columbian tooth
to cf. Albertosaurus.
References-
Buckley, McCrea and Currie, 2005. Theropod teeth from the Upper
Cretaceous Kaskapau (Middle Turonian) and the Wapiti (Upper Campanian -
Lower Maastrichtian) formations of north-eastern British Columbia,
Canada. Journal of Vertebrate Paleontology. 25(3), 40A-41A.
Fanti and Miyashita, 2009. A high latitude vertebrate fossil assemblage from
the Late Cretaceous of west-central Alberta, Canada: Evidence for dinosaur nesting
and vertebrate latitudinal gradient. Palaeogeography, Palaeoclimatology, Palaeoecology.
275, 37-53.
Fanti, Currie and Burns, 2015. Taphonomy, age, and paleoecological implication
of a new Pachyrhinosaurus (Dinosauria: Ceratopsidae) bonebed from the
Upper Cretaceous (Campanian) Wapiti Formation of Alberta, Canada. Canadian Journal
of Earth Sciences. 52(4), 250-260.
Reid, 2016. A review of dinosaurian body fossils from British Columbia, Canada. PeerJ Preprints. 4:e1369v3.
undescribed Tyrannosauridae (Russell, 1935)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
Material- (CMN coll.) teeth (37.5 mm) (Russell, 1935)
(GSC 8724; = CMN 8724) tooth (Ford and Chure, 2001)
(MR-4:74) tooth (Baszio, 1997)
(RTMP 20021) (juvenile) tooth (Ryan and Russell, 2001)
References- Russell, 1935. Fauna of the Upper Milk River beds, southern
Alberta. Transactions of the Royal Society of Canada, series 3, section 4. 29,
115-127.
Baszio, 1997. Investigations on Canadian dinosaurs: Systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ford and Chure, 2001. Ghost lineages and the paleogeographic and temporal distribution
of tyrannosaurids. Journal of Vertebrate Paleontology. 21(3), 50A-51A.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). In Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press. 279-297.
undescribed Tyrannosauridae (Lambe, 1902)
Middle-Late Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
Material- (CMN 57080) quadratojugal (Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017)
(CMN coll.) several teeth (to 90 mm), metatarsal fragments, several
phalanges, unguals (Lambe, 1902)
(YPM 9834) (YPM online)
Comments- Carr et al. (2017)
note "an isolated tyrannosaurid quadratojugal from the upper Campanian
strata of southern Alberta (CMN 57080)" has a lateral quadratojugal
foramen as in juvenile Tyrannosaurus and the oldest known Daspletosaurus "horneri" (MOR 1130).
Reference- Lambe, 1902. New genera and species from the Belly River Series
(Mid-Cretaceous). Geological Survey of Canada Contributions to Canadian Palaeontology.
3(2), 25-81.
Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017. A new tyrannosaur
with evidence for anagenesis and crocodile-like facial sensory system.
Scientific Reports. 7:44942.
undescribed Tyrannosauridae (Ryan and Russell, 2001)
Middle Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
Material- (RTMP 88.86.4) tooth (Ryan and Russell, 2001)
(RTMP 96.62.71-90; within that range of numbers) (juvenile and adult) fifteen teeth (Peng, Russell and Brinkman, 2001)
(RTMP 96.62.48 or 49) (juvenile) two premaxillary teeth (Peng, Russell and Brinkman, 2001)
(RTMP coll.) tooth (Cullen, Fanti, Capobianco, Ryan and Evans, 2016)
(?UC coll.) tooth (Frampton, 2006)
Comments- These may be referrable to the co-occuring "Thanatotheristes degrootorum".
Ryan and Russell (2001) list RTMP 88.86.4 as a reference
Tyrannosauridae indet. specimen from the Foremost Formation,
"uncatalogued TMP teeth" as a reference Aublysodon
sp. specimen, and list additional teeth as representing both. The
uncatalogued and additional teeth include those described by Peng et
al. (2001) from three localities in the Foremost Formation (five plus
two unserrated 'Aublysodon'
teeth from PHR-1; nine from PHR-2 and one from SPS). They state
"they are very similar in morphology to those from other upper
Cretaceous strata. The incompleteness of the material makes precise
assignment at lower taxonomic level impossible."
Cullen et al. (2016) listed a single tyrannosaurid tooth from the PK microsite.
Cullen and Evans (2016) list one tyrannosaurid tooth from the PHRN site, which equates with Frampton's (2006) thesis.
References- Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages
(exclusive of mammals) from the Foremost and Oldman Formations of the
Judith River Group (Campanian) of southeastern Alberta: An illustrated
guide. Provincial Museum of Alberta Natural History Occasional Paper. 25, 54 pp.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press.
279-297.
Brinkman, Russell, Eberth and Peng, 2004. Vertebrate palaeocommunities
of the lower Judith River Group (Campanian) of southeastern Alberta,
Canada, as interpreted from vertebrate microfossil assemblages.
Palaeoecology, Palaeoclimatology, Palaeoecology. 213, 295-313.
Frampton, 2006. Taphonomy and palaeoecology of mixed
invertebrate-vertebrate fossil assemblage in the Foremost Formation
(Cretaceous, Campanian), Milk River Valley, Alberta. Masters thesis, University
of Calgary. 294+ pp.
Cullen and Evans, 2016. Palaeoenvironmental drivers of vertebrate
community composition in the Belly River Group (Campanian) of Alberta,
Canada, with implications for dinosaur biogeography. BMC Ecology. 16,
52.
Cullen, Fanti, Capobianco, Ryan and Evans, 2016. A vertebrate microsite
from a marine-terrestrial transition in the Foremost Formation
(Campanian) of Alberta, Canada, and the use of faunal assemblage data
as a paleoenvironmental indicator. Palaeoecology, Palaeoclimatology,
Palaeoecology. 444, 101-114.
undescribed Tyrannosauridae (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
Material-
(RTMP 92.30.219) tooth (Ryan and Russell, 2001)
(RTMP 96.62.48) (juvenile) tooth (Ryan and Russell, 2001)
(RTMP coll.) twelve teeth (Chiba et al., 2015)
(YPM-PU 24515) (YPM online)
Comments- These may belong to Gorgosaurus or Daspletosaurus.
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press.
279-297.
Chiba, Ryan, Braman, Eberth, Scott, Brown, Kobayashi and Evans, 2015. Taphonomy
of a monodominant Centrosaurus apertus (Dinosauria: Ceratopsia) bonebed
from the upper Oldman Formation of southeastern Alberta. Palaios. 30, 655-667.
undescribed Tyrannosauridae (Molnar and Carpenter, 1989)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
Material- (AMNH 3838) several teeth (AMNH online as Allosaurus? sp.)
(CMN 12) quadrate (Carr, 1996)
(CMN 16) anterior dentary (Carr, 1996)
(CMN 23) partial dentary (Carr, 1996)
(CMN 116a) (juvenile) tooth (Molnar and Carpenter, 1989)
(CMN 947) quadrate condyle (Carr, 1996)
(CMN 1822) (juvenile) tooth (Molnar and Carpenter, 1989)
(CMN 2196; see also CMN 2196 under Albertosaurus) surangular (Carr, 1996)
(CMN 2225) dentaries (Carr, 1996)
(CMN 2248) anterior dentary (Carr, 1996)
(CMN 2637) partial premaxilla (Carr, 1996)
(CMN 41104) (juvenile) premaxillary tooth (Currie, Rigby and Sloan, 1990)
(FMNH PR864) anterior dentary (Carr, 1996)
(FMNH PR1196) anterior dentary (Carr, 1996)
(ROM 43296) lacrimal (Carr, 1996)
(RTMP 66.31.93) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 79.10.59) (juvenile) tooth (9.8 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 80.8.192) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 80.16.864) tooth (80 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 80.16.1202) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 81.16.197) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 81.19.79) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 81.19.263) (juvenile) tooth (15.5 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.367) (juvenile) premaxillary tooth (Currie, Rigby and Sloan, 1990;
Ryan and Russell, 2001)
(RTMP 82.20.457) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 85.6.134) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 86.77.122) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 86.77.123) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 87.36.81) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 87.46.24) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 88.4.7) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP coll.) 44 teeth (Ryan, Russell, Eberth and Currie, 2001)
(RTMP coll.) (juvenile) five teeth (Ryan, Russell, Eberth and Currie, 2001)
Comments- These are probably from Gorgosaurus libratus or Daspletosaurus
sp. nov..
References- Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian,
Montana, U.S.A.) referred to the genus Aublysodon. Geobios. 22, 445-454.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press. 107-125.
Carr, 1996. Tyrannosauridae (Dinosauria: Theropoda) from the Dinosaur Park Formation
(Judith River Group, Upper Cretaceous: Campanian) of Alberta. Masters Thesis.
University of Toronto. 358 pp.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). In Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press. 279-297.
Ryan, Russell, Eberth and Currie, 2001. The taphonomy of a Centrosaurus
(Ornithischia: Certopsidae) bone bed from the Dinosaur Park Formation (Upper
Campanian), Alberta, Canada, with comments on cranial ontogeny. Palaios. 16(5),
482-506.
undescribed Tyrannosauridae (Langston, 1975)
Maastrichtian, Late Cretaceous
St. Mary River Formation, Alberta, Canada
Material- (CMN 9589) tooth
(CMN 9723) tooth
(CMN 10650) tooth
(CMN 10651) tooth
(CMN 10652) tooth
(CMN 10675) tooth
Reference- Langston, 1975. The ceratopsian dinosaurs and associated lower
vertebrates from the St. Mary River Formation (Maestrichtian) at Scabby Butte,
southern Alberta. Canadian Journal of Earth Sciences. 12(9), 1576-1608.
undescribed Tyrannosauridae (Russell, 1930)
Maastrichtian, Late Cretaceous
Lower Ravenscrag Formation, Saskatchewan, Canada
Material- teeth
Reference- Russell, 1930. Upper Cretaceous dinosaur faunas of North America.
Proceedings of the American Philosophical Society. 69, 133-159.
undescribed Tyrannosauridae (Mongelli and Varricchio, 1998)
Early Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- (MOR 414) five teeth (MOR online)
(MOR 1116) tibia, metatarsal (MOR online)
(same as MOR 414?) teeth (Mongelli and Varricchio, 1998)
Reference- Mongelli and Varricchio, 1998. Theropod teeth of the Lower
Two Medicine Formation (Campanian) of northwestern Montana. Journal of Vertebrate
Paleontology. 18(3), 65A.
undescribed Tyrannosauridae (Redman, Moore and Varricchio, 2015)
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
Material-
(MOR 268) (~715 mm; embryo) (skull ~89 mm) dentary (29 mm) (Funston,
Powers, Whitebone, Brusatte, Scannella, Horner and Currie,
2020)
(MOR 313) tibiae (MOR online)
(MOR 586) quadratojugal, quadrate (MOR online)
(Old Trail Museum coll.; = MOR 953) cranial elements (MOR online)
(YPM-PU 24967) (YPM online)
teeth (Redman, Moore and Varricchio, 2015)
Comments- Funston et al. (2020)
state embryonic dentary MOR 268 "already exhibits distinctive
tyrannosaurine characters", so it is probably Daspletosaurus, two
species of which seem to be present in the formation.
References- Redman, Moore and Varricchio, 2015. A new vertebrate microfossil
locality in the Upper Two Medicine Formation in the vicinity of Egg Mountain.
Journal of Vertebrate Paleontology. Program and Abstracts 2015, 201-202.
Funston, Powers, Whitebone, Brusatte, Scannella, Horner and Currie,
2020. Baby tyrannosaur bones from the Late Cretaceous of western North
America. The Society of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 147-148.
undescribed Tyrannosauridae (Carrano, 1998)
Late Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
Material- (MOR 468) cranial fragment, vertebrae, pelvic element, femur,
phalanx (MOR online)
(MOR 553E) femur (912 mm) (Carrano, 1998)
(MOR 553E-6-19-91-69) radius (MOR online)
(MOR 565) tooth (MOR online)
(MOR 589) braincase (MOR online)
Reference- Carrano, 1998. The evolution of dinosaur locomotion: Functional
morphology, biomechanics, and modern analogs. PhD Thesis, The University of
Chicago. 424 pp.
undescribed Tyrannosauridae (Sahni, 1972)
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Material- (AMNH 8515) anterior dentary tooth (Sahni, 1972)
(MOR 028) tooth (MOR online)
(MOR 033) teeth (MOR online)
(MOR 034) premaxillary teeth (MOR online)
(MOR 395) maxilla (MOR online)
(MOR 644) cranial fragments (MOR online)
(MOR 657) partial skull and skeleton including maxilla, metatarsal II, phalanx
II-1, phalanx II-2, ungual II, metatarsal III (528 mm), phalanx III-1, phalanx
III-2, phalanx III-3, ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2,
phalanx IV-3, phalanx IV-4, metatarsal V (Carrano, 1998)
(MOR 769) partial skeleton (MOR online)
(MOR 1029) tooth (MOR online)
(MOR 1061) fragmentary nasal (MOR online)
(RMDRC coll.) thirteenth dorsal vertebra, sacrum (~600 mm) (Maltese, 2017 online)
(UCMP 136555) tooth (UCMP online)
(UCMP 136556) tooth (UCMP online)
(YPM-PU 21545) (YPM online)
(YPM-PU 21848) (YPM online)
(YPM-PU 22250) (YPM online)
(YPM-PU 22339) (YPM online)
(YPM-PU 22402) (YPM online)
(YPM-PU 23476) (YPM online)
(YPM-PU 24965) (YPM online)
(YPM-PU 24971) (YPM online)
References- Sahni, 1972. The vertebrate fauna of the Judith River Formation,
Montana. Bulletin of the AMNH. 147(6), 412 pp.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Maltese, 2017 online. Digging a tyrannobutt. RMDRC paleo lab. 7-5-2017.
undescribed tyrannosaurid (MOR online)
Late Maastrichtian, Late Cretaceous
Livingston Formation, Montana, US
Material- (MOR 002) postcranial skeleton (MOR online)
undescribed Tyrannosauridae (Carpenter, 1982)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (MOR 064) tooth fragment (MOR online)
(MOR 072) tooth fragment (MOR online)
(MOR 336) centrum (MOR online)
(RTMP 87.112.33) (juvenile) tooth (Molnar and Carpenter, 1989)
(RTMP 87.114.7) (juvenile) tooth (Molnar and Carpenter, 1989)
(UCMP 124367) (juvenile) tooth (6.1 mm) (Carpenter, 1982)
(UCMP 88125, 109015, 109018, 119676, 119677, 120135, 120262, 120306, 120352,
123373, 123508, 123509, 123545, 124486, 12549) teeth (Ford and Chure, 2001)
(UCMP 112003) distal tibia (Ford and Chure, 2001)
(UCMP 119508) phalanges (Ford and Chure, 2001)
(UCMP 119578, 119678, 120080, 120137) tooth fragments (Ford and Chure, 2001)
(UCMP 119579, 119580, 120017, 120048) phalanges (Ford and Chure, 2001)
(UCMP 119725) metatarsal? (Ford and Chure, 2001)
(UCMP 119785) six vertebrae (Ford and Chure, 2001)
(UCMP 119786) two teeth (Ford and Chure, 2001)
(UCMP 119787) two teeth (Ford and Chure, 2001)
(UCMP 119788) three teeth (Ford and Chure, 2001)
(UCMP 119853) (juvenile) four teeth (Ford and Chure, 2001)
(UCMP 119854) two tooth fragments (Ford and Chure, 2001)
(UCMP 119929) seven teeth (Ford and Chure, 2001)
(UCMP 119931) four teeth (Ford and Chure, 2001)
(UCMP 119932) tooth (Ford and Chure, 2001)
(UCMP 119933) tooth (Ford and Chure, 2001)
(UCMP 119934) six tooth fragments (Ford and Chure, 2001)
(UCMP 119935) three caudals. (Ford and Chure, 2001)
(UCMP 119936) five phalanges (Ford and Chure, 2001)
(UCMP 120001) two unguals (Ford and Chure, 2001)
(UCMP 120081) ungual (Ford and Chure, 2001)
(UCMP 120136) two teeth (Ford and Chure, 2001)
(UCMP 120194) three tooth fragments (Ford and Chure, 2001)
(UCMP 120260) two teeth (Ford and Chure, 2001)
(UCMP 120261) six tooth fragments (Ford and Chure, 2001)
(UCMP 120263) caudal vertebra (Ford and Chure, 2001)
(UCMP 120305) two teeth (Ford and Chure, 2001)
(UCMP 120307) pedal phalanx. (Ford and Chure, 2001)
(UCMP 120339) two teeth (Ford and Chure, 2001)
(UCMP 120340) ungual (Ford and Chure, 2001)
(UCMP 120341) caudal vertebra (Ford and Chure, 2001)
(UCMP 120843) two teeth (Ford and Chure, 2001)
(UCMP 120844) two manual phalanges (Ford and Chure, 2001)
(UCMP 120845) four ungual fragments (Ford and Chure, 2001)
(UCMP 123342) eight teeth (Ford and Chure, 2001)
(UCMP 123343) 20+ teeth (Ford and Chure, 2001)
(UCMP 123344) seven teeth (Ford and Chure, 2001)
(UCMP 123345) 20+ teeth (Ford and Chure, 2001)
(UCMP 123346) 40+ teeth (Ford and Chure, 2001)
(UCMP 123347) metatarsal (Ford and Chure, 2001)
(UCMP 123567) three teeth (Ford and Chure, 2001)
(UCMP 124484, cast) tooth (Ford and Chure, 2001)
(UCMP 124485) four teeth (Ford and Chure, 2001)
(YPM 54459) (YPM online)
(YPM 54461) (YPM online)
(YPM 55508) (YPM online)
(YPM 55519) (YPM online)
(YPM 55532) (YPM online)
(YPM 55540) (YPM online)
(YPM 55541) (YPM online)
(YPM 55559) (YPM online)
(YPM 55569) (YPM online)
(YPM 55597) (YPM online)
(YPM 55618) (YPM online)
teeth, bones (Triebold, 1997)
Comments- These are probably Tyrannosaurus, based on their provenance.
References- Carpenter, 1982. Baby dinosaurs from the Late Cretaceous
Lance and Hell Creek formations and a description of a new species of theropod.
Contributions to Geology, University of Wyoming. 20(2), 123-134.
Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian, Montana, U.S.A.)
referred to the genus Aublysodon. Geobios. 22, 445-454.
Triebold, 1997. The Sandy Site: Small dinosaurs from the Hell Creek Formation
of South Dakota. In Wolberg, Stump and Rosenberg (eds.). Dinofest International.
245-248.
Ford and Chure, 2001. Ghost lineages and the paleogeographic and temporal distribution
of tyrannosaurids. Journal of Vertebrate Paleontology. 21(3), 50A-51A.
undescribed tyrannosaurid (Hoganson, Erickson and Getman, 1994)
Maastrichtian, Late Cretaceous
Timber Lake Member of Fox Hills Formation, North Dakota, US
Material- tooth
Reference- Hoganson, Erickson and Getman, 1994. Reptiles of the Timber
Lake Member (Cretaceous: Maastrichtian), Fox Hills Formation, North Dakota.
Journal of Vertebrate Paleontology. 14(3), 29A.
undescribed tyrannosaurid (Young, 1987)
Late Campanian, Late Cretaceous
Williams Fork Formation of Mesaverde Group, Colorado, US
Material- tooth
Reference- Young, 1987. Remains of ancient life in Cretaceous rocks of
the Dinosaur Triangle. Dinosaur Triangle Paleontological Field Trip, 1987. 45-54.
undescribed tyrannosaurid (Molnar and Carpenter, 1989)
Late Maastrichtian, Late Cretaceous
Denver Formation, Colorado, US
Material- (UCMP 38060) (juvenile) tooth
Comments- This is probably Tyrannosaurus, based on its provenance.
Reference- Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian,
Montana, U.S.A.) referred to the genus Aublysodon. Geobios. 22, 445-454.
undescribed Tyrannosauridae (Kirkland, Lucas and Estep, 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah, US
Material- (OMNH 23635) tooth (Parrish, 1999)
(OMNH 24309; in part) (juvenile) tooth (Parrish, 1999)
Comments- Parrish (1999) listed OMNH 24635 as Tyrannosauridae and 24309
as cf. Aublysodon.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. In Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian)
of southern Utah. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
undescribed tyrannosaurid (Parker and Rowley, 1989)
Early Campanian, Late Cretaceous
Blackhawk Formation, Utah, US
Material- skull (Madsen pers. comm. 1985 to Parker and Rowley, 1989)
tooth (Robison pers. comm. 1984 to Parker and Rowley, 1989)
Comments- Parker and Rowley (1989) referred the skull to Albertosaurus
sp., citing a Parker and Balsley in prep. publication which never emerged.
Reference- Parker and Rowley, 1989. Dinosaur footprints from a coal mine
in east-central Utah. In Gillette and Lockley (ed.). Dinosaur Tracks and Traces.
Cambridge University Press. 360-366.
unnamed tyrannosaurid (Thomson and Irmis, 2010)
Late Campanian, Late Cretaceous
Neslen Formation of Mesaverde Group, Utah, US
Material- (UNMH VP 16395) incomplete fibula, partial metatarsal II, metatarsal
IV (490 mm), partial metatarsal V
Comments- Thomson and Irmis (2010) stated this specimen is similar to
Daspletosaurus torosus
in having a slender ridge along the posterior surface of metatarsal IV
proximal to the distal metatarsal III attachment site. Thomson et
al. (2013) believed this was more derived than Appalachiosaurus
but outside derived tyrannosaurines, most similar to albertosaurines and Daspletosaurus.
References- Thomson and Irmis, 2010. First occurence of a tyrannosaurid
(Dinosauria, Theropoda) from the Neslen Formation (Late Cretaceous), Book Cliffs
area, Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2010,
175A.
Thomson, Irmis and Loewen, 2013. First occurrence of a tyrannosaurid
dinosaur from the Mesaverde Group (Neslen Formation) of Utah: Implications for
Upper Campanian Laramidian biogeography. Cretaceous Research. 43, 70-79.
undescribed Tyrannosauridae (Parrish, 1999)
Late Campanian, Late Cretaceous
Kaiparowits Formation, Utah, US
Material- (MNA HM-6; in part) tooth (Parrish, 1999)
(OMNH 21960) tooth (Parrish, 1999)
(OMNH 21961) tooth (Parrish, 1999)
(RAM 8395) (subadult) partial dentary (Zanno et al., 2013)
(RAM 9132) femur, partial tibia, astragalus, calcaneum, partial metatarsal III,
phalanx III-1, phalanx III-2 (Lofgren, Cripe and Everhart, 2007)
(UCM 8304) tooth (Parrish, 1999)
(UCM 8323?; not 83239 as listed) tooth (Parrish, 1999)
(UCM 8626) tooth (Parrish, 1999)
(UCM 8642; in part) tooth (Parrish, 1999)
(UCM 8647) tooth (Parrish, 1999)
(UCM 8659) tooth (Parrish, 1999)
(UCM 8671) tooth (Parrish, 1999)
(UMNH VP 11302) (adult) tooth, caudal vertebra, femur, tibia, fibula, metatarsal
III, pedal phalanx, ungual (Zanno et al., 2013)
(UMNH VP 12223) humerus (Zanno et al., 2013)
(UMNH VP 12586) (juvenile) cranial material including partial frontal, parietals
and partial dentary (Zanno et al., 2013)
(UMNH VP 16161) teeth, partial limb elements (Zanno et al., 2013)
(UMNH VP 16225) parietals (Zanno et al., 2013)
(UMNH VP 16692) fragmentary limb elements, pedal phalanx, ungual (Zanno et al.,
2013)
(UMNH VP 16693) skull fragments including partial dentary, limb fragments including
pedal phalanx and ungual (Zanno et al., 2013)
(RAM or UMNH VP coll.) (adult) lacrimal (Zanno et al., 2013)
Comments- These may belong to Teratophoneus, the only named tyrannosauroid
from this formation.
References- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian)
of southern Utah. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013. Late Cretaceous theropod
dinosaurs of southern Utah. In Titus and Loewen (eds.). At the Top of the Grand
Staircase: The Late Cretaceous of Southern Utah. Indiana University Press. 504-525.
undescribed Tyrannosauridae (Gilmore, 1946)
Late Maastrichtian, Late Cretaceous
North Horn Formation, Utah, US
Material- teeth, manual ungual, distal metatarsal II or IV
Comments- These were identified as deinodontids, representing both large
and small individuals. It's possible some belong to Tyrannosaurus, which
is known from the formation, or other coelurosaurs.
Reference- Gilmore, 1946. Reptilian fauna of the North Horn Formation
of central Utah. United States Geological Survey Professional Paper. 210-C,
29-53.
undescribed Tyrannosauridae (Konecny, 1994)
Fort Crittenden Formation, Arizona, US
Late Campanian, Late Cretaceous
Material- (Konecny private coll.) two tooth fragments (Konecny, 1994)
(UALP 1925; = A 25) tooth (McCord, 1997)
(UALP 1927; = A 26) tooth (McCord, 1997)
(UALP 1928; = A 27) tooth (McCord, 1997)
References- Konecny, 1994. Dinosaurs of Arizona. MAPS Digest. 17(4),
17-24.
McCord, 1997. An Arizona titanosaurid sauropod and revision of the Late Cretaceous
Adobe Canyon fauna. Journal of Vertebrate Paleontology. 17(3), 620-622.
Krzyzanowski, Lucas and Heckert, 2001. Late Campanian (Judithian) vertebrate
fauna of the Fort Crittenden Formation, Southeastern Arizona. Journal of Vertebrate
Paleontology. 21(3), 69A-70A.
undescribed Tyrannosauridae (Lucas, Basabilvazo and Lawton, 1990)
Campanian, Late Cretaceous
Ringbone Formation, New Mexico, US
Material- (NMMNH P-3050) proximal caudal centrum
(NMMNH P-12997) tooth
Reference- Lucas, Basabilvazo and Lawton, 1990. Late Cretaceous dinosaurs
from the Ringbone Formation, southwestern New Mexico, U.S.A. Cretaceous Research.
11, 343-349.
undescribed Tyrannosauridae (Armstrong-Ziegler, 1980)
Late Campanian, Late Cretaceous
Fruitland Formation, New Mexico, US
Material- (KUVP 85370) limb element (Carr and Williamson, 2000)
(MNA Pl.1623) three teeth (Armstrong-Ziegler, 1980)
(NMMNH P-30077) tooth (Carr and Williamson, 2000)
(NMMNH P-32590) tooth (Carr and Williamson, 2000)
(PMA P 73.30.1) metatarsus (480 mm) (Holtz, 1994)
References- Armstrong-Ziegler, 1980. Amphibia and Reptilia from the Campanian
of New Mexico. Fieldiana, Geology. 4, 39 pp.
Holtz, 1994. The arctometatarsalian pes, an unusual structure of the metatarsus
of Cretaceous Theropoda (Dinosauria: Saurischia). Journal of Vertebrate Paleontology.
14(4), 480-519.
Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria: Coelurosauria)
from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico
Museum of Natural History and Science. Bulletin. 17, 113-146.
undescribed Tyrannosauridae (Carr and Williamson, 2000)
Late Campanian, Late Cretaceous
Fruitland or Lower Kirtland Formation, New Mexico, US
Material- (LACM 45985) tooth fragments
(NMMNH P-22693) partial pedal phalanx
(NMMNH P-22908) distal metatarsal II, distal metatarsal III
(NMMNH P-27744) distal metatarsal IV, fragments
(NMMNH P-27773) digit I
(NMMNH P-27787) proximal rib
(NMMNH coll.) 19+ teeth [see Carr and Williamson, 2000 for numbers]
(USNM 365551) incomplete pubis, femur (665 mm), tibia, metatarsal
Reference- Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria:
Coelurosauria) from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New
Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 113-146.
undescribed Tyrannosauridae (Brown, 1910)
Late Campanian-Maastrichtian, Late Cretaceous
Fruitland or Kirtland Formation, New Mexico, US
Material- (AMNH 2479 in part; syntype of Dysganus encaustus) tooth
(Carr and Williamson, 2000)
(AMNH coll.) teeth ()
(KUVP 14958) mandibular fragment (Carr and Williamson, 2000)
(KUVP 15135) tooth (Carr and Williamson, 2000)
(KUVP 15145) teeth (Carr and Williamson, 2000)
(KUVP 15234-15235) two unguals (Carr and Williamson, 2000)
(KUVP 15262) tooth (Carr and Williamson, 2000)
(KUVP 15605) teeth (Carr and Williamson, 2000)
(KUVP 16042) tooth, phalanx (Carr and Williamson, 2000)
(KUVP 96846) femur (Carr and Williamson, 2000)
(KUVP 96861) astragalus (Carr and Williamson, 2000)
(KUVP 96878) tooth (Carr and Williamson, 2000)
(KUVP 96888) mandible (Carr and Williamson, 2000)
(PMU.R35; = R1235?) anterior dentary (Sullivan and Williamson, 1997)
(PMU.R36; = R56?) tooth (Sullivan and Williamson, 1997)
(PMU.R85) partial dentary (Carr and Williamson, 2000)
(USNM 10754) metatarsus (540 mm) (Holtz, 1994)
(USNM coll.) teeth (Gilmore, 1916)
(uncollected) (~8 m) vertebrae, limb elements (Brown, 1910)
References- Brown, 1910. The Cretaceous Ojo Alamo beds of New Mexico
with description of the new dinosaur genus Kritosaurus. Bulletin of the
American Museum of Natural History. 28, 267-284.
Brown, 1914. Cretaceous Eocene correlations in New Mexico, Wyoming, Montana.
Bulletin of the Geological Society of America. 25, 355-380.
Gilmore, 1916. Vertebrate faunas of the Ojo Alamo, Kirtland and Fruitland formations.
United States Geological Survey, Professional Paper. 98Q, 279-308.
Holtz, 1994. The arctometatarsalian pes, an unusual structure of the metatarsus
of Cretaceous Theropoda (Dinosauria: Saurischia). Journal of Vertebrate Paleontology.
14(4), 480-519.
Sullivan and Williamson, 1997. Additions and corrections to Sternberg’s
San Juan Basin Collection, Paleontological Museum, University of Uppsala, Sweden.
New Mexico Geological Society Guidebook, 48th Field Conference, Mesozoic Geology
and Paleontology of the Four Corners Region, 1997. 255-257.
Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria: Coelurosauria)
from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico
Museum of Natural History and Science Bulletin. 17, 113-146.
undescribed tyrannosaurid (Molnar and Carpenter, 1989)
Late Campanian-Maastrichtian, Late Cretaceous
Kirtland Formation, New Mexico, US
Material- (KU 12419) (juvenile) tooth (Molnar and Carpenter, 1989)
(PMU.R40; =R1240?) pedal ungual (Sullivan and Williamson, 1997)
References- Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian,
Montana, U.S.A.) referred to the genus Aublysodon. Geobios. 22, 445-454.
Sullivan and Williamson, 1997. Additions and corrections to Sternberg’s
San Juan Basin Collection, Paleontological Museum, University of Uppsala, Sweden.
New Mexico Geological Society Guidebook, 48th Field Conference, Mesozoic Geology
and Paleontology of the Four Corners Region, 1997. 255-257.
undescribed Tyrannosauridae (Lehman and Carpenter, 1990)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-22976; = UNM B-828?) femur (995 mm) (Lehman and Carpenter,
1990)
(NMMNH P-25073) proximal scapula (Carr and Williamson, 2000)
(NMMNH P-27281) pedal phalanx (Carr and Williamson, 2000)
(NMMNH P-27620) pedal phalanges (Carr and Williamson, 2000)
(NMMNH P-29164) tibia (Carr and Williamson, 2000)
(NMMNH P-30072) partial pedal phalanx (Carr and Williamson, 2000)
(NMMNH P-30074) caudal vertebra (Carr and Williamson, 2000)
(NMMNH P-30075) pedal phalanx fragment (Carr and Williamson, 2000)
(NMMNH coll.) twelve teeth [see Carr and Williamson, 2000 for numbers]
References- Lehman and Carpenter, 1990. A partial skeleton of the tyrannosaurid
dinosaur Aublysodon from the Upper Cretaceous of New Mexico. Journal
of Paleontology. 64, 1026-1032.
Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria: Coelurosauria)
from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico
Museum of Natural History and Science Bulletin. 17, 113-146.
undescribed Tyrannosauridae (Gilmore, 1916)
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
Material- (NMMNH 12999, others; =UNM FKK-077, 078, 079, 080) four teeth
(Lucas et al., 1987)
(NMMNH P-20879) pedal phalanx (Hunt and Lucas, 1992)
(NMMNH P-25071) distal femur (Carr and Williamson, 2000)
(NMMNH P-25085) tibia (993 mm) (Carr and Williamson, 2000)
(NMMNH P-26276) manual ungual (Carr and Williamson, 2000)
(NMMNH P-27276) distal metatarsal II (Carr and Williamson, 2000)
(NMMNH P-27280) tooth (16.6x8.7x3.8 mm) (Williamson and Brusatte, 2014)
(NMMNH P-27287) two caudal centra, neural arch (Carr and Williamson, 2000)
(NMMNH P-27461) partial dorsal vertebra (Carr and Williamson, 2000)
(NMMNH P-28923) distal caudal vertebra (Carr and Williamson, 2000)
(NMMNH P-28926) partial phalanx (Carr and Williamson, 2000)
(NMMNH P-30014) two distal caudal vertebrae (Carr and Williamson, 2000)
(NMMNH P-33903a) tooth (?x?x6.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33903b) tooth (?x~8.5x5.1 mm) (Williamson and Brusatte, 2014)
(NMMNH coll.) >64 teeth [see Carr and Williamson, 2000 for numbers]
(USNM 8346) dentary (Gilmore, 1916)
(USNM 8355) (juvenile) premaxillary tooth (Gilmore, 1916)
References- Gilmore, 1916. Vertebrate faunas of the Ojo Alamo, Kirtland
and Fruitland formations. United States Geological Survey, Professional Paper.
98Q, 279-308.
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.
Gilmore, 1935. On the Reptilia of the Kirtland Formation of New Mexico, with
descriptions of new species of fossil turtles. Proceedings of the United States
National Museum. 83(2978), 159-188.
Lucas et al., 1987. Dinosaurs, the age of the Fruitland and Kirtland Formations,
and the Cretaceous-Tertiary boundary in the San Juan Basin, New Mexico. In Fassett
and Rigby (eds.). The Cretaceous-Tertiary Boundary in the San Juan and Raton
Basins, New Mexico and Colorado. Geological Society of America Special Paper.
209, 35-50.
Hunt and Lucas, 1992. Stratigraphy, paleontology and age of the Fruitland and
Fruitland Formations (Upper Cretaceous), San Juan basin, New Mexico. New Mexico
Geological Society Guidebook. 43, 217-239.
Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria: Coelurosauria)
from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico
Museum of Natural History and Science. Bulletin. 17, 113-146.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
undescribed Tyrannosauridae (Lehman, 1982)
Late Maastrichtian, Late Cretaceous
Tornillo Group, Texas, US
Material- (TAMU 1372) tooth
(TMM 31201-1) tooth
(TMM 31201-6) tooth
(TMM 31221-1) femur
(TMM 40573-1) tibia
(TMM 41395-3) tooth
(TMM 41541-2) tooth
(TMM 41541-3) tooth
(TMM 42291-1) tooth
(TMM 42291-2) tooth
(TMM 49710-1) tooth
(TMM AM 144) tooth
Reference- Lehman, 1982. A ceratopsian bone bed from the Aguja Formation
(Upper Cretaceous) Big Bend National Park, Texas. Masters thesis, University
of Texas at Austin. 210 pp.
undescribed Tyrannosauridae (Lehman, 1985)
Late Maastrichtian, Late Cretaceous
Javelina Formation, Texas, US
Reference- Lehman, 1985. Stratigraphy, sedimentology, and paleontology of
Upper Cretaceous (Campanian-Maastrichtian) sedimentary rocks in Trans-Pecos
Texas. PhD thesis, University of Texas at Austin. 299 pp.
undescribed Tyrannosauridae (Lehman, 1985)
Late Maastrichtian, Late Cretaceous
El Picacho Formation, Texas, US
Reference- Lehman, 1985. Stratigraphy, sedimentology, and paleontology of
Upper Cretaceous (Campanian-Maastrichtian) sedimentary rocks in Trans-Pecos
Texas. PhD thesis, University of Texas at Austin. 299 pp.
undescribed Tyrannosauridae (Westgate, Brown and Pittman, 2002)
Campanian, Late Cretaceous
San Carlos Formation, Mexico
Reference- Westgate, Brown and Pittman, 2002. Discovery of dinosaur remains
in coastal deposits near Ojinaga, Mexico. Journal of Vertebrate Paleontology.
22(3), 118A-119A.
undescribed Tyrannosauridae (Murry, Boyd, Wolleben and Wilson, 1960)
Late Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Material- teeth (Rivera-Sylva, Frey, Stinnesbeck, Padilla Gutierrez, Gonzalez
Gonzalez and Amezcua Torres, 2015)
Reference- Murry, Boyd, Wolleben and Wilson, 1960. Late Cretaceous fossil
locality, eastern Parras Basin, Coahuila, Mexico. Journal of Paleontology. 34(2),
368-370.
Rivera-Sylva, Frey, Stinnesbeck, Padilla Gutierrez, Gonzalez Gonzalez and Amezcua
Torres, 2015. The Late Cretaceous Las Aguilas dinosaur graveyard, Coahuila,
Mexico. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 203.
undescribed Tyrannosauridae (Molnar, 1974)
Late Campanian, Late Cretaceous
El Gallo Formation, Mexico
Material- (AMNH 7755) tooth, three vertebrae, partial femur (1.165 m) (Carrano,
1998)
(IGM 4302; = LACM 20886) (juvenile) premaxillary tooth (Molnar, 1974)
(IGM coll.; = LACM 7253/28999) (juvenile) tooth (Ford and Chure, 2002)
(IGM coll.; = LACM 3294/24580) (juvenile) tooth (Ford and Chure, 2002)
(LACM 17715 in part) three teeth (Morris, 1981)
(LACM 28237) metatarsal (Molnar, 1974)
teeth and bone fragments (Morris, 1981)
teeth (Rodriguez de la Rosa and Aranda-Manteca, 1999)
metatarsal IV, pedal phalanx, phalangeal fragments (Peecook, Wilson, Silson,
Hernandez and Montellano-Ballesteros, 2010)
Comments- Molnar (1974) reported only teeth and a metatarsal were known
from the El Gallo Formation, the latter of which is longer and more gracile
than Labocania. Hernandez-Rivera (1997) noted cf. Albertosaurus
remains from the El Gallo Formation, which are probably the same material (e.g.
Morris, 1981 noting carnosaur teeth and bone fragments "as large as Gorgosaurus
or Tyrannosaurus"). The IGM teeth are laterally compressed and they
are serrated on both carinae. Denticles are chisel-shaped, decrease in size
toward the base and tip of the tooth, and the tyrannosaurid blood grooves run
obliquely from between the denticles and extend toward the tooth base.
References- Molnar, 1974. A distinctive theropod dinosaur from the Upper
Cretaceous of Baja California (Mexico). Journal of Paleontology. 48(5), 1009-1017.
Morris, 1981. A new species of hadrosaurian dinosaur from the Upper Cretaceous
of Baja California ?Lambeosaurus laticaudus. Journal of Paleontology.
55(2), 453-462.
Hernandez-Rivera, 1997. Mexican dinosaurs. In Currie and Padian (eds.). Encyclopedia
of Dinosaurs. Academic Press. 433-437.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Rodriguez-de la Rosa and Aranda-Manteca, 1999. Theropod teeth from the Late
Cretaceous El Gallo Formation, Baja California, Mexico. VII International Symposium
on Mesozoic Terrestrial Ecosystems, Buenos Aires, Abstracts. 56.
Ford and Chure, 2002. "Aublysodon" teeth from the El Gallo
Formation (Late Campanian) of Baja California: The southernmost record of tyrannosauroid
theropods. Mesa Southwest Museum Bulletin. 8, 75-89.
Peecook, Wilson, Silson, Hernandez and Montellano-Ballesteros, 2010. New tyrannosauroid
remains from the Late Cretaceous 'El Gallo' Formation of Baja de California,
Mexico. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 144A.
undescribed Tyrannosauridae (Lucas, Kues and Gonzalez-Leon, 1995)
Late Campanian-Maastrichtian, Late Cretaceous
Corral de Enmedio Formation, Mexico
Material- (IRGNM-210) tooth
(IRGNM-211) partial hindlimb including partial tibia, fibula, phalanges
(IRGNM coll.) teeth, elements
Reference- Lucas, Kues and Gonzalez-Leon, 1995. Paleontology of the Late
Cretaceous Cabullona Group, northeastern Sonora. Geological Society of America,
Special paper. 301, 143-165.
Albertosaurinae Currie, Hurum and Sabath,
2003
Definition- (Albertosaurus sarcophagus <- Tyrannosaurus
rex) (Holtz, 2004; modified from Currie et al., 2003)
= Albertosaurini Olshevsky, 1995
Diagnosis- (after Carr, 2005) elongate lacrimal pneumatic recess; postorbital
boss does not approach dorsal margin of bone; postorbital boss position adjacent
to orbit; posterodorsal margin of posterior postorbital process is concave;
posterior process of postorbital stops short of posterior margin of laterotemporal
fenestra; medial margin of joint surface for the quadratojugal on the quadrate
extends vertically; ceiling of basisphenoid recess is inflated; dorsal process
of palatine is short; dorsal process of palatine is anteroposteriorly elongate;
dorsal process of palatine extended vertically; the dorsal margin of the lateral
cnemial process extends anteroventrally at a steep angle; shaft of pedal phalanx
I-1 is wide; ventrally, the joint surface of the lateral condyle of pedal phalanx
I-1 reaches or extends past the posterior margin of the collateral ligament
pit; the ventral margin of the proximal surface of pedal phalanx IV-2 is trilobate;
in dorsal view the distal condyle of pedal phalanx IV-2 extends into the supracondylar
pit.
Comments- This is generally thought to include Gorgosaurus libratus
and perhaps Appalachiosaurus, in addition to Albertosaurus sarcophagus.
Olshevsky (1995) created the tribe Albertosaurini as a paraphyletic taxon including
not only Albertosaurus and Gorgosaurus, but Daspletosaurus
as well.
References- Olshevsky, 1995. The origin and evolution of the tyrannosaurids.
Kyoryugaku Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.
Currie, Hurum and Sabath, 2003. Skull structure and evolution in tyrannosaurid
dinosaurs. Acta Palaeontologica Polonica. 48(2), 227-234.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis, University of Toronto. 1170 pp.
Gorgosaurus Lambe, 1914
G. libratus Lambe, 1914
= Deinodon libratus (Lambe, 1914) Matthew and Brown, 1922
= Gorgosaurus “sternbergi” Matthew and Brown, 1922
= Gorgosaurus sternbergi Matthew and Brown, 1923
= Deinodon sternbergi (Matthew and Brown, 1923) Kuhn, 1965
= Albertosaurus libratus (Lambe, 1914) Russell, 1970
= Albertosaurus sternbergi (Matthew and Brown, 1923) Russell, 1970
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (CMN 2120) (8.25 m; adult) skull (985 mm), mandible (950 mm),
partial third cervical vertebra, partial fourth cervical vertebra, partial fifth
cervical vertebra, partial sixth cervical vertebra, partial seventh cervical
vertebra, partial eighth cervical vertebra, partial ninth cervical vertebra,
partial tenth cervical vertebra, six cervical ribs, first dorsal vertebra, second
dorsal vertebra (93 mm), third dorsal vertebra (97 mm), fourth dorsal vertebra
(100 mm), fifth dorsal vertebra (102 mm), sixth dorsal vertebra, seventh dorsal
vertebra, eight dorsal vertebra, ninth dorsal vertebra, tenth dorsal vertebra,
eleventh dorsal vertebra (134 mm), twelfth dorsal vertebra (150 mm), twenty
dorsal ribs, fused anterior gastralia, eighteen gastralia, (sacrum 690 mm) first
sacral vertebra (138 mm), second sacral vertebra (128 mm), third sacral vertebra
(130 mm), fourth sacral vertebra (134 mm), fifth sacral vertebra (160 mm), first
caudal vertebra, second caudal vertebra, third caudal vertebra (159 mm), fourth
caudal vertebra (144 mm), fifth caudal vertebra (143 mm), sixth caudal vertebra
(162 mm), seventh caudal vertebra (126 mm), eighth caudal vertebra (144 mm),
ninth caudal vertebra (117 mm), tenth caudal vertebra (144 mm), eleventh caudal
vertebra (140 mm), twelfth caudal vertebra (142 mm), thirteenth caudal vertebra
(162 mm), fourteenth caudal vertebra (149 mm), fifteenth caudal vertebra (139
mm), sixteenth caudal vertebra (134 mm), seventeenth caudal vertebra (132 mm),
eighteenth caudal vertebra (123 mm), nineteenth caudal vertebra (122 mm), twentieth
caudal vertebra (117 mm), twenty-first caudal vertebra (113 mm), twenty-second
caudal vertebra (108 mm), twenty-third caudal vertebra (104 mm), twenty-fourth
caudal vertebra (104 mm), partial twenty-sixth caudal vertebra, twenty-seventh
caudal vertebra (81 mm), twenty-eighth caudal vertebra (72 mm), chevrons 2-22,
scapula (876 mm), coracoid (210 mm), humerus (324 mm), radius (156 mm), ulna
(180 mm), radiale, ulnare, intermedium, distal carpal I, distal carpal II, metacarpal
I (48 mm), phalanx I-1 (98 mm), manual ungual I (82 norm, 95 mm adc), metacarpal
II (98 mm), phalanx II-1 (57 mm), phalanx II-2 (83 mm), manual ungual II (64
mm), metacarpal III (64 mm), partial ilium (984 mm), pubis (980 mm), ischium
(762 mm), femur (1.04 m), tibia (1 m), fibula (883 mm), astragalus (208 wide,
300 mm tall), calcaneum, distal tarsal II, distal tarsal III, distal tarsal
IV, metatarsal I (115 mm), phalanx I-1 (100 mm), pedal ungual I (95 mm), metatarsal
II (508 mm), phalanx II-1 (164 mm), phalanx II-2 (121 mm), pedal ungual II (120
mm), metatarsal III (594 mm), phalanx III-1 (163 mm), phalanx III-2 (122 mm),
phalanx III-3 (93 mm), pedal ungual III (45 mm), metatarsal IV (546 mm), phalanx
IV-1 (110 mm), phalanx IV-2 (92 mm), phalanx IV-3 (65 mm), phalanx IV-4 (50
mm), pedal ungual IV (104 mm), metatarsal V (216 mm)
Referred- (AMNH 5358) skull, skeleton (Currie and Russell, 2005)
(AMNH 5423) anterior maxilla, skull roof, dentary, skeleton including four caudal
vertebrae, pelvic fragments, hindlimb including femur (600 mm), tibia (630 mm)
and metatarsus (440 mm) (Russell, 1970)
(AMNH 5428) dorsal rib fragments
....(USNM 12814; =AMNH 5428) (1.01 tons; 18 year old adult) skull (795 mm),
mandible including dentary, cervical series, cervical ribs, dorsal series, dorsal
ribs, gastralia, sacrum, first caudal neural spine, caudal vertebrae 17-19,
scapulocoracoid, ilium, pubis, proximal ischium, femora (880 mm), tibiae (850
mm), fibulae, metatarsi (535 mm), pedes (Matthew and Brown, 1923)
(AMNH 5432) (1.28 tons; 22 year old adult) fragmentary skull (anterior maxilla,
fragmentary nasals, jugals, fragmentary braincase; partial ectopterygoid), partial
coronoid, four caudal vertebrae, pelvic fragments, hindlimb including tibia
(910 mm), astragalus, metatarsus (590 mm) and phalanges (Russell, 1970)
(AMNH 5434; = AMNH 5336 of Matthew and Brown, 1923 and Russell, 1970) (adult)
skull (1.05 m), mandible (1.025 m), cervical vertebrae, dorsal vertebrae, sacrum,
scapulocoracoid (965 mm), humerus (328 mm), radius (163 mm), ulna (200 mm),
metacarpal I (60 mm), phalanx I-1 (145 mm), metacarpal II (110 mm), hindlimb
excluding pes (femur ~1.093 m) (Matthew and Brown, 1923)
(AMNH 5458) (8.6 m, 2.5 tons, adult) skull (990 mm), mandible (985 mm), incomplete
skeleton including presacral column (2.55 m total), sacrum (665 mm), caudal
vertebrae 1-3, caudal vertebrae 20-30, scapulcaoracoid, ilium (1.04 m), partial
pubis, partial ischium, femur (1.025 m), tibia (990 mm), metatarsus (625 mm),
phalanx III-1 (173 mm) (Matthew and Brown, 1923)
(AMNH 5664; holotype of Gorgosaurus sternbergi) (5.8 m, 700 kg, juvenile)
incomplete skull (678 mm), mandible (690 mm), nine cervical vertebrae (600 mm
total), cervical ribs, thirteen dorsal vertebrae (1.042 mm total), dorsal ribs,
gastralia, sacrum (472 mm), caudal vertebrae 1-24 (2.45 m total), chevrons 1-24,
scapulocoracoid (620 mm), humeri (205 mm), radius (100 mm), ulnae (125 mm),
metacarpal I (40 mm), metacarpal II (60 mm), ilium (695 mm), pubis (610 mm),
ischium (465 mm), femur (700 mm), tibia (748 mm), fibula (680 mm), metatarsus
(480 mm), phalanx III-1 (78 mm) (Matthew and Brown, 1922)
(CMN 2193) (adult) surangular (Russell, 1970)
(CMN 2250) ilium (Russell, 1970)
(CMN 2270) (juvenile) maxilla (Russell, 1970)
(CMN 8782) fragmentary skull, incomplete manus, hindlimb fragments, incomplete
pes (Russell, 1970)
(CMN 11593) proximal tail, pelvis, hindlimbs including femur (940 mm), tibia
(900 mm) and metatarsus (580 mm), skin impressions (Russell, 1970)
(CMN 11814) braincase (Russell, 1970)
(CMN 12063) (juvenile) maxilla (Russell, 1970)
(ROM 436) (juvenile) partial premaxilla, maxillary fragment (Russell, 1970)
(ROM 683) premaxillae, incomplete maxillae, nasal fragment, anterior dentary
(Russell, 1970)
(ROM 4591) (adult) nasals (Russell, 1970)
(ROM 1237) skeleton lacking presacral vertebrae and one hindlimb (Russell, 1970)
(ROM 1247) (juvenile) skull (lacking premaxillae, postorbitals, palatines, pterygoids)
(750 mm), mandible (lacking coronoids), skeleton including femur (730 mm), tibia
(775 mm), metatarsus (542 mm) (Russell, 1970)
(ROM 1422) (adult) partial skull (premaxilla, incomplete maxillae, incomplete
nasals, lacrimal, postorbital, partial squamosal, quadratojugal, quadrates,
partial palatine, partial pterygoid), partial surangular, angular (Russell,
1970)
(RTMP 67.9.164) dentary (445 mm) (Currie, 2003b)
(RTMP 68.3.1) pelvis, hindlimbs (Currie, 2003b)
(RTMP 73.30.1) (750 kg; 14 year old subadult) proximal tail, pelvis, hindlimb
including tibia (805 mm), metatarsus (515 mm) (femur ~804 mm) (Currie, 2003b)
(RTMP 80.16.485) (juvenile) frontal, tooth (Molnar and Carpenter, 1989)
(RTMP 83.36.100) (juvenile) skull (Carr, 1999)
(RTMP 83.36.134) dentary (Currie, 2003b)
(RTMP 85.11.3) (juvenile) maxilla (Currie, Rigby and Sloan, 1990)
(RTMP 86.144.1) (4.5 m; 230 kg; 7 year old juvenile) skull (500 mm), dentary,
splenial, prearticular, skeleton including femur (542 mm) (Carr, 1999)
(RTMP 91.36.500) (5.1 m; juvenile) incomplete skeleton including skull (670
mm), mandible, furcula (172 mm), humerus, fibula, phalanx II-2, metatarsal III
(460 mm) and pedal ungual III (Makovicky and Currie, 1998)
(RTMP 91.163.1) skull, skeleton (Currie, 2003b)
(RTMP 94.12.155) (3 m; juvenile) cranial fragments (~364 mm), mandibles (Carr,
1999)
(RTMP 94.12.602) (10 m; 1.12 tons; 18 year old adult) skeleton including skull,
stapes, dorsal ribs, gastralia, furcula (225 mm), femur (916 mm) and fibula
(Makovicky and Currie, 1998)
(RTMP 94.143.1) (5.8 m; 496 kg; 10 year old subadult) skull (620 mm), mandible,
two cervical vertebrae, four cervical ribs, five dorsal vertebrae, dorsal neural
spine, seven dorsal ribs (two fragmentary), gastralia, ten caudal vertebrae,
five chevrons, vertebral fragment, partial ilium, femoral fragment (~626 mm)
(Tanke and Currie, 2000)
(RTMP 95.5.1) skull, dentary, skeleton (Currie, 2003b)
(RTMP 99.33.1) (607 kg; 14 year old subadult) skull, dentary, skeleton including
femur (750 mm) (Currie, 2003b)
(RTMP 99.55.170) dentary (Currie, 2003b)
(RTMP 2000.12.11) skull (Currie, 2003b)
(UALVP 10) (adult) skull (900 mm), mandibles including dentary, fifth through
eighth cervical vertebrae, six dorsal vertebrae, ribs, gastralia, partial forelimb
including humerus, partial hindlimbs including metatarsus and phalanx III-1
(Russell, 1970)
(UALVP 49500) (juvenile) complete skeleton (Bradley and Currie, 2013)
fragmentary skeleton (Currie and Russell, 2005)
Middle Campanian, Late Cretaceous
Oldman Formation, Alberta, Canada
(FMNH PR2211) (130 kg; 5 year old juvenile) postcranial skeleton including ribs,
gastralia, femur (445 mm), and fibula (Currie, 2003b)
(RTMP 2005.12.117) incomplete skull including braincase (Ali et al., 2008)
Middle-Late Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
(RTMP 67.14.3) frontal (Currie, 2003b)
(RTMP 80.16.924) frontal, parietal (Ali et al., 2008)
(RTMP 81.39.8) frontal (Currie, 2003b)
(RTMP 82.16.181) frontal (Currie, 2003b)
(RTMP 82.28.1) cranial element, dentary, vertebra, gastralia and pedal ungual (Currie, 2003b)
(RTMP 86.49.29) dentary (Currie, 2003b)
(RTMP 91.36.533) frontal, parietal (Currie, 2003b)
(RTMP 92.36.76) frontal, parietal (Currie, 2003b)
(RTMP 92.36.82) squamosal (Currie, 2003b)
(RTMP 92.36.749) dentary (Currie, 2003b)
Middle Campanian, Late Cretaceous
Oldman Formation, Saskatchewan, Canada
material (Tokaryk, 1988)
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
(AMNH 3963) (21 years old) dentary, premaxillary tooth (Cope, 1876)
(MOR 28) tooth (MOR online)
(MOR 33) ten teeth (MOR online)
(MOR 34) three premaxillary teeth (MOR online)
(MOR 644) skull fragments (MOR online)
(MOR 657) partial maxillae, skull fragments, teeth, four vertebrae, neural arch,
rib fragments, femora, tibiae, fibulae, metatarsal II, metatarsal III, metatarsal
IV, metatarsal fragments, fourteen phalanges, five pedal unguals, fragments
(MOR online)
(MOR 769) premaxillary tooth fragment, two neural arches, three ribs, chevron,
pubis, ischium, fragments (MOR online)
(Peebles coll.) incomplete skeleton (Potera, 1995)
(private coll.) (at least four individuals) material including femora (328 mm
to 1.132 m) (Currie and Eberth, 2010)
Late Cretaceous
North America
(ROM 672) specimen including metacarpal I, metacarpal II, metacarpal III, femur
(795 mm), tibia (813 mm) and metatarsal III (520.5 mm) (Smith and Galton, 1990)
(ROM 762) scapulocoracoid (Parsons and Parsons, 2009)
(ROM 1246) dentary (Currie, 2003b)
(ROM 3520) frontal (Currie, 2003b)
(TCM 2001.89.1) maxilla (568 mm), lacrimal, dentary (580 mm), scapula (675 mm),
humerus (305 mm), ulna (180 mm), ilium (865 mm), femur (825 mm), metatarsal
II (490 mm), metatarsal IV (500 mm) (Larson, 2008)
Diagnosis- (after Carr, 2005) lacrimal pneumatic recess dorsoventrally
deep; laterosphenoid extends dorsomedially to contact the parietal; the distal
joint surface of pedal phalanx II-2 does not reach the anterior margin of either
collateral ligament pit; in medial view the proximal joint surface of pedal
phalanx II-3 does not extend onto the dorsal surface of the bone.
Comments- Pharris (DML 1996) suggested the name "Albertogorgon lambei"
for several specimens (AMNH 5336, AMNH 5664, FMNH PR308, ROM 1247 and USNM 12814)
based on differences Paul (1988) noted between these and G. libratus
(CMN 2120, AMNH 5458, RTMP 85.62.1). However, the name is unpublished, FMNH
PR308 is Daspletosaurus, sternbergi has priority as the species,
and recent studies (Carr, 1996; Currie et al., 2003; Carr, 2005) have not recognized
the reality of such a taxon. Indeed, in Carr's (2005) specimen-level analysis,
AMNH 5664 was in a clade with the holotype (CMN 2120), separate from AMNH 5336,
5458 and ROM 1247. This completely mixes "Albertogorgon" and Gorgosaurus
specimens, showing the division is artificial.
AMNH 5434 was described by Matthew and Brown (1923) as AMNH 5336, which was
repeated in the literature by Russell and others. AMNH 5336 is actually a Daspletosaurus
specimen which was called AMNH 5434 by Matthew and Brown, and later moved to
the FMNH as PR308. MOR 557 is listed by Currie (2003b) as G. libratus,
but is on the MOR website as a Tyrannosaurus specimen.
At at talk in Chicago after the Armour Symposium, Currie (2001) reported skin
impressions associated with the holotype of Gorgosaurus, which lacked
scales. Bell et al. (2017) noted "during restoration of the specimen in
the 1980's; however, these were later covered over with plaster and the moulds
taken of the skin could not be relocated at the time of writing." Some
other specimens from Dinosaur Park show this same morphology. Tanke (DML 1996)
reported a small patch of skin associated with a partial tyrannosaurid skeleton
(vertebrae, dorsal ribs, gastralia, ilium impression, limb bones impressions,
astragalus) from Alberta presumably stored in the RTMP. The tyrannosaurid was
~8-9 m long, and the skin impression (though associated with a gastralium and
ilial impression) could not be placed anywhere specifically on the body due
to the skeleton's disarticulation. It preserved small reticulate scales similar
to hadrosaurids. Tanke also reported tyrannosaurid skin impressions on MOR specimens.
Most recently, Bell et al. described a skin impression on CMN 11593
References- Cope, 1876. Descriptions of some vertebrate remains from
the Fort Union Beds of Montana. Proceedings of the Academy of Natural Sciences
of Philadelphia. 28, 248-261.
Lambe, 1914. On a new genus and species of carnivorous dinosaur from the Belly
River Formation of Alberta, with a description of the skull of Stephanosaurus
marginatus from the same horizon. Ottawa Naturalist. 28, 13-20.
Lambe, 1917. The Cretaceous theropodous dinosaur Gorgosaurus. Geological
Survey of Canada, Memoir. 100, 84 pp.
Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus
from the Cretaceous of Alberta. American Museum of Natural History Bulletin.
46, 367-385.
Matthew and Brown, 1923. Preliminary notices of skeletons and skulls of Deinodontidae
from the Cretaceous of Alberta. American Museum Novitates. 89, 10 pp.
Kuhn, 1965. Saurischia (Supplementum 1). In Fossilium Catalogus 1. Animalia.
109, 94 pp.
Russell, 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National
Museum of Natural Sciences, Publications in Palaeontology. 1, 1-34.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Tokaryk, 1988. Preliminary vertebrate faunal list of the Oldman Formation, Saskatchewan.
Journal of Vertebrate Paleontology. 8(3), 28A.
Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian,
Montana, U.S.A.) referred to the genus Aublysodon. Geobios. 22, 445-454.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press. 107-125.
Smith and Galton, 1990. Osteology of Archaeornithomimus asiaticus (Upper
Cretaceous, Iren Dabasu Formation, People's Republic of China). Journal of Vertebrate
Paleontology. 10(2), 255-265.
Potera, 1995. Amateur fossil hunters dig up trouble in Montana. Science. 268(5208), 198-199.
Carr, 1996. Tyrannosauridae (Dinosauria: Theropoda) from the Dinosaur Park Formation
(Judith River Group, Upper Cretaceous: Campanian) of Alberta. MS Thesis. University
of Toronto. 358 pp.
Pharris, DML 1996. https://web.archive.org/web/20201113234528/http://dml.cmnh.org/1996Jan/msg00171.html
Tanke, DML 1996. https://web.archive.org/web/20191009075258/http://dml.cmnh.org/1996Feb/msg00497.html
Carr, 1998. Tyrannosaurid (Dinosauria: Theropoda) craniofacial ontogeny: Comparative
parsimony analysis of ontogenetic characters. Journal of Vertebrate Paleontology.
18(3), 31A.
Makovicky and Currie, 1998. The presence of a furcula in tyrannosaurid theropods,
and its phylogenetic and functional implications. Journal of Vertebrate Paleontology.
18(1), 143-149.
Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria).
Journal of Vertebrate Paleontology. 19(3), 497-520.
Tanke and Currie, 2000. Head-biting behavior in theropod dinosaurs: Paleobathological
evidence. Gaia. 15, 167-184.
Currie, 2001. Feathered dinosaurs and the origin of birds. Talk in Chicago on
5-13.
Currie, 2003a. Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from
the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences.
40(4), 651-665.
Currie, 2003b. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous
of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Erickson, Makovicky, Currie, Norell, Yerby and Brochu, 2004. Gigantism and comparative
life-history parameters of tyrannosaurid dinosaurs. Nature. 430, 772-775.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Currie and Russell, 2005. The geographic and stratigraphic distribution of articulated
and associated dinosaur remains. In Currie and Koppelhus (eds.). Dinosaur Provincial
Park, a spectacular ecosystem revealed. Indiana University Press. 537-569.
Ali, Zelenitsky, Therrien and Weishampel, 2008. Homology of the "ethmoid
complex" of tyrannosaurids and its implications for the reconstruction
of the olfactory apparatus of non-avian theropods. Journal of Vertebrate Paleontology.
28(1), 123-133.
Bell, 2008. A medley of maladies: Multiple paleopathologies in a specimen of
Gorgosaurus libratus (Tyrannosauridae). Journal of Vertebrate Paleontology.
28(3), 50A.
Larson, 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In
Larson and Carpenter (eds.). Tyrannosaurus rex: The Tyrant King. Indiana
University Press. 102-128.
Parsons and Parsons, 2009. Further descriptions of the osteology of Deinonychus
antirrhopus (Saurischia, Theropoda). Bulletin of the Buffalo Society of
Natural Sciences. 38, 43-54.
Buckley, Larson, Reichel and Samman, 2010. Quantifying tooth variation within
a single population of Albertosaurus sarcophagus (Theropoda: Tyrannosauridae)
and implications for identifying isolated teeth of tyrannosaurids. Canadian
Journal of Earth Sciences. 47, 1227-1251.
Currie and Eberth, 2010. On gregarious behavior in Albertosaurus. Canadian
Journal of Earth Sciences. 47(9), 1277-1289.
Bradley and Currie, 2013. Tooth and postcranial growth rates in a juvenile Gorgosaurus
libratus. Journal of Vertebrate Paleontology. Program and Abstracts 2013,
92.
Bradley and Currie, 2014. Gregariousness in Gorgosaurus: Studying social
behaviour in non-avian theropod dinosaurs (Dinosauria: Saurischia). Journal
of Vertebrate Paleontology. Program and Abstracts 2014, 96.
Bradley, Glasier and Currie, 2015. Comparing tooth macrowear in a juvenile and
adult specimen of Gorgosaurus libratus: Changes in feeding behavior throughout
ontogeny in tyrannosaurids. Journal of Vertebrate Paleontology. Program and
Abstracts 2015, 95.
Bell, Campione, Persons, Currie, Larson, Tanke and Bakker, 2017. Tyrannosauroid
integument reveals conflicting patterns of gigantism and feather evolution.
Biology Letters. 13: 20170092.
Voris, 2018. Cranial anatomy and ontogeny of Gorgosaurus libratus (Tyrannosauridae: Albertosaurinae). Masters thesis, University of Calgary. [pp]
"Albertosaurus" incrassatus
(Cope, 1876) Huene, 1932
= Laelaps incrassatus Cope, 1876
= Dryptosaurus incrassatus (Cope, 1876) Hay, 1902
= Deinodon incrassatus (Cope, 1876) Osborn, 1902
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Syntypes- (AMNH 3962) (subadult) first maxillary tooth (25 mm)
.... (juvenile) first maxillary tooth (14 mm)
Comments- The type teeth (AMNH 3962) are distinctive in being labiolingually
wider than mesiodistally long (13.5 mm vs. 12 mm for the larger specimen; 8
mm vs. 6 mm for the smaller), and unlike tyrannosaurid premaxillary teeth, the
distal carina is median in position. The mesial carina twists lingually at its
base and both carinae are serrated. These characters are only known in maxillary
tooth 1 of Gorgosaurus, which is present in the same strata further north.
While Lambe (1904) indicates the first dentary tooth of Albertosaurus
is wider than long as well, he also notes this tooth is D-shaped as in premaxillary
teeth. The size indicates they are young specimens.
Cope (1876) later referred a dentary and a premaxillary tooth (AMNH 3963) to
this species. He describes one or two anterior teeth (but not the first dentary
tooth, which is D-shaped as in Albertosaurus) as being transversely uncompressed
or even expanded, and lists measurements for the second tooth as having a labiolingual
diameter of 18 mm and a mesiodistal diameter of 13 mm. Cope's measurement may
not have taken into account the rotation of the carinae in tyrannosaurid anterior
dentary teeth though, as Tyrannosaurus has a second dentary tooth only
65% as wide labiolingially as mesiodistally, but it is rotated to appear transversely
broader than long in dorsal view of the dentary. Currie (2003) later referred
this specimen to Gorgosaurus.
Cope (1892) described two specimens (CMN 5600 and 5601) from the Horseshoe Canyon
Formation of Alberta as further specimens of Laelaps incrassatus. Lambe
(1903, 1904) published more detailed descriptions of these specimens as Dryptosaurus
incrassatus. The combination was first used by Hay (1902) because Laelaps
was found to be preoccupied and replaced with Dryptosaurus by Marsh.
However, Lambe's (1904) statement that Cope's original teeth and dentary are
more likely Deinodon, making the Horseshoe Canyon specimens the types
of incrassatus is incorrect. The name must stick with Cope's original
holotype teeth. Osborn (1905) recognized this and created the taxon Albertosaurus
sarcophagus for the Horseshoe Canyon specimens, to distinguish them from
the Judith River type material of incrassatus.
References- Cope, 1876. Descriptions of some vertebrate remains from
the Fort Union Beds of Montana. Paleontological Bulletin. 22, 1-14.
Cope, 1876. Descriptions of some vertebrate remains from the Fort Union Beds
of Montana. Proceedings of the Academy of Natural Sciences of Philadelphia.
28, 248-261.
Cope, 1892. Skull of the dinosaurian Laelaps incrassatus Cope. Proceedings
of the American Philosophical Society. 30, 240-245.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 1-868.
Osborn, 1902. On Vertebrata of the Mid-Cretaceous of the Northwest Territory.
I: Distinctive characters of the Mid-Cretaceous fauna. Contributions to Canadian
Palaeontology. 3, 1-21.
Lambe, 1903. The lower jaw of Dryptosaurus incrassatus (Cope). The Ottawa
Naturalist. 175, 133-139.
Lambe, 1904. On Dryptosaurus incrassatus (Cope), from the Edmonton Series
of the North West Territory. Geological Survey of Canada Contributions to Canadian
Palaeontology. 3(3), 1-27.
Osborn, 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs.
Bulletin of the American Museum of Natural History. 21, 259-265.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous
of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Albertosaurus Osborn, 1905
A. sarcophagus Osborn, 1905
= Deinodon sarcophagus (Osborn, 1905) Matthew and Brown, 1922
= Albertosaurus arctunguis Parks, 1928
= Deinodon arctunguis (Parks, 1928) Kuhn, 1939
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
Holotype- (CMN 5600) partial skull, incomplete mandibles (970 mm), limb
elements and other postcrania (lost)
Paratype- (CMN 5601) partial skull (maxilla- 457 mm), incomplete mandibles,
neural spine, sacral neural arches, partial ilium, distal tibia, astragalus
(248 mm wide), metatarsal IV (505 mm), three pedal unguals (109 mm)
Referred- (AMNH 5218 small individual) (~4.1 m) specimen including femur
(954 mm), tibia (850 mm), metatarsus (515 mm) and pedal phalanx III-3 (42 mm)
(Russell and Chamney, 1967)
(AMNH 5218) (multiple individuals, including an 18 and 20 year old) two dentaries,
fourteen vertebrae, two chevrons, scapula, coracoid, two humeri, pubes, three
femora, four tibiae, fibula, two astragali, calcaneum, metatarsal I, four phalanges
I-1, pedal ungual I, metatarsal II, four phalanges II-1, four phalanges II-2,
three phalanges III-1, four phalanges III-2, five phalanges III-3 (99 mm), metatarsal
IV, six phalanges IV-1, phalanx IV-2, four phalanges IV-3, phalanx IV-4, five
metatarsals, five pedal phalanges, six pedal unguals (Currie, 2000)
(AMNH 5222) incomplete skull (Russell and Chamney, 1967)
(AMNH 5224) (several individuals) includes skull, tail and hindlimbs (Ford and
Chure, 2001)
(AMNH 5228; listed as II and III by Eberth and Currie, 2010) (~6.9 m, 17 year
old) metatarsal III, metatarsal IV (465 mm) (Currie, 2000)
(AMNH 5229) (~5.1 m, 8 year old juvenile) metatarsal II, metatarsal IV (394
mm) (Currie, 2000)
(AMNH 5230; lost) metatarsus (Currie, 2000)
(AMNH 5231; listed as astragalus, mts II-IV and 4 phalanges by Eberth and Currie,
2010) (~7.6 m) astragalus, two distal tarsals, metatarsal II, phalanx II-1,
phalanx II-2, metatarsal IV (510 mm), phalanx IV-2 (Currie, 2000)
(AMNH 5232) (~8.1 m) two distal tarsals, metatarsal II, metatarsal III (560
mm), metatarsal IV (521 mm), metatarsal V, two phalanges, pedal ungual (Currie,
2000)
....(AMNH 5226) 25 caudal vertebrae (Currie, 2000)
....(AMNH 5227) tibia (815 mm), fibula, astragalus (Carrano, 1998)
(AMNH 5233; = AMNH 5223 of Erickson et al., 2010?, which is an Anodontosaurus
specimen) (~6.0 m, 11 year old) metatarsal II, phalanx II-1, metatarsal III
(480 mm), phalanx III-1, metatarsal IV (426 mm) (Currie, 2000)
(AMNH 5234) (~6.8 m, 15 year old) astragalus, metatarsal II, metatarsal III
(440 mm), phalanx III-1, metatarsal IV (452 mm), phalanx IV-1 (Currie, 2000)
(AMNH 5235) (~7.3 m, 17 year old) femur (870 mm), tibia, metatarsal II, metatarsal
III (510 mm), metatarsal IV (486 mm), ungual (Currie, 2000)
?(AMNH 5255) hindlimb including femur (692 mm) and proximal tibia (Osborn, 1916)
(CMN 2196; = CMN 1912-5?; see also CMN 2196 under Dinosaur Park Fm. indet.)
gastralia, scapulocoracoid (Lambe, 1914)
(CMN 11315; previously referred to Daspletosaurus) (juvenile) cranial
elements, gastralia, scapula (470 mm), coracoid (110 mm), furcula (162 mm),
humerus (225 mm), radius (96 mm), ulna (120 mm), metacarpal I (32 mm), phalanx
I-1 (63 mm), manual ungual I (57 mm), metacarpal II (58 mm), phalanx II-1 (28
mm), phalanx II-2 (~40 mm), manual ungual II (61 mm), metacarpal III (38 mm),
ilium (~675 mm), pubis (~600 mm), ischium (488 mm), femur (665 mm), tibia (736
mm), astragalus, metatarsus (448 mm) (Russell and Chamney, 1967)
(?Perth Museum coll.; lost) several vertebrae, partial scapulocoracoid, humeri,
femur, tibia (Sternberg, 1915)
(ROM 807; holotype of Albertosaurus arctunguis) (8.6 m, 2.5 tons) tenth
dorsal vertebra (110 mm), eleventh dorsal vertebra (120 mm), twelfth dorsal
vertebra (125 mm), thirteenth dorsal vertebra (130 mm), four fragmentary anterior
dorsal ribs, seventh to twelfth dorsal ribs (465-840 mm), fused anterior gastralia(?),
gastralia, sacrum (675 mm), first caudal vertebra (130 mm), second caudal vertebra,
third caudal vertebra (133 mm), first chevron, scapula (740 mm), coracoid (148
mm), humerus (303 mm), radius (136 mm), ulna (163 mm), radiale, intermedium,
ulnare, metacarpal I (40 mm), phalanx I-1 (85 mm), manual ungual I (115 mm on
curve), metacarpal II (80 mm), phalanx II-1 (45 mm), phalanx II-2 (70 mm), manual
ungual II (100 mm on curve), ilium (980 mm), pubis (1.03 m), ischium (660 mm),
femur (1.02 m), tibia (980 m), fibula (875 mm), astragalus (260 mm wide), calcaneum,
distal tarsal IV, metatarsal II (540 mm), metatarsal III (590 mm), phalanx III-1
(200 mm), phalanx III-2 (140 mm), phalanx III-3 (100 mm), metatarsal IV (558
mm), metatarsal V (225 mm) (Parks, 1928)
(ROM 12790) occipital condyle (Carr 1999)
(RTMP 81.9.1) skeleton missing caudal series (Olshevsky, 1995)
(RTMP 81.10.1) (8 m; 1.14 tons; 24 year old adult) (skull ~970 mm) maxilla,
jugal, ectopterygoid, quadrates, surangular, angular, prearticular, partial
postcrania missing tail (femur- 895mm, tibia- 970 mm, metatarsus- 610 mm) (Coy,
1982)
....(Stefanuk coll.) pedal phalanx II-1 (Tanke and Currie, 2010)
(RTMP 81.31.59) premaxillary tooth (Erickson, 1995)
(RTMP 82.13.3) jugal, postorbital (Carr, 2010)
(RTMP 82.13.30) femur (902.0 mm) (Carrano, 1998)
(RTMP 85.98.1) (subadult) incomplete skull, mandibles, incomplete skeleton including
three anterior cervical vertebrae, five dorsal ribs, gastralia, astragalus,
metatarsal III and metatarsal V (Maier, 1985)
(RTMP 86.64.1) (6.5 m; 760 kg; 15 year old subadult) incomplete skeleton including
skull, mandible, atlas, axis, third cervical vertebra, incomplete fourth cervical
vertebra, vertebral column, ribs, gastralia, chevron, scapulocoracoid, furcula
(192 mm), forelimb including humerus (212 mm), pubes, ischium, partial femur
(~782 mm), tibia (750 mm), fibula, astragalus, metatarsals (mtIII 482 mm) and
pedal phalanges (Carrano, 1998)
(RTMP 86.205.1) (adult) quadrate, basioccipital, braincase elements, cranial
fragments, mandibles, cervical vertebrae, ribs, gastralia, scapulae, forelimb
elements, ilium, pubis, hindlimb elements including metatarsals and phalanges
(Danis, 1986)
(RTMP 89.17.15) partial pterygoid (Bell and Currie, 2014)
?...(RTMP 89.17.53) (~8-8.5 m) incomplete maxilla (Bell, 2007; described in
Bell and Currie, 2014)
?...(UALVP 52743) incomplete dentary (Bell and Currie, 2014)
(RTMP 94.25.6) dentary (390 mm) (Currie, 2003)
(RTMP 94.186.1) vertebral fragments, dorsal rib fragments, gastralial fragments,
hindlimb fragments including astragalus (width ~200 mm), skin impressions (Currie,
2003)
(RTMP 95.25.83) maxilla (Currie, 2003)
(RTMP 95.91.1) jugal (Carr, 2010)
(RTMP 97.58.1) maxilla (lost), dentaries (lost), ribs, gastralia, about eight
distal caudal vertebrae, tibia (lost) (Currie, 2003)
(RTMP 98.59.4) rib fragment (Bell, 2010)
(RTMP 98.59.5) rib fragment (Bell, 2010)
(RTMP 98.63.1) pedal ungual (120 mm) (Eberth and Currie, 2010)
(RTMP 98.63.2) palatine (Eberth and Currie, 2010)
(RTMP 98.63.3) tooth (47 mm) (Eberth and Currie, 2010)
(RTMP 98.63.6) metacarpal (Eberth and Currie, 2010)
(RTMP 98.63.7) gastralia (Eberth and Currie, 2010)
(RTMP 98.63.9) rib fragment (Eberth and Currie, 2010)
(RTMP 98.63.10) tooth (38 mm) (Eberth and Currie, 2010)
(RTMP 98.63.11) anterior dentary tooth (28 mm) (Reichel, 2010)
(RTMP 98.63.12) tooth (Eberth and Currie, 2010)
(RTMP 98.63.14) tooth (27 mm) (Eberth and Currie, 2010)
(RTMP 98.63.15) tooth (61 mm) (Eberth and Currie, 2010)
(RTMP 98.63.16) tooth (62 mm) (Eberth and Currie, 2010)
(RTMP 98.63.19) tooth (65 mm) (Eberth and Currie, 2010)
(RTMP 98.63.20) tooth (~30 mm) (Eberth and Currie, 2010)
(RTMP 98.63.21) tooth (Eberth and Currie, 2010)
(RTMP 98.63.22) tooth (65 mm) (Eberth and Currie, 2010)
(RTMP 98.63.23) tooth (40 mm) (Eberth and Currie, 2010)
(RTMP 98.63.25) distal caudal vertebra (41 mm) (Eberth and Currie, 2010)
(RTMP 98.63.26) tooth (42 mm) (Eberth and Currie, 2010)
(RTMP 98.63.27) tooth (12 mm) (Eberth and Currie, 2010)
(RTMP 98.63.29) tooth (31 mm) (Eberth and Currie, 2010)
(RTMP 98.63.30) lateral gastralium (Eberth and Currie, 2010)
(RTMP 98.63.31) tooth (Eberth and Currie, 2010)
(RTMP 98.63.32) tooth (Eberth and Currie, 2010)
(RTMP 98.63.34) premaxillary tooth (Eberth and Currie, 2010)
(RTMP 98.63.35) tooth (Eberth and Currie, 2010)
(RTMP 98.63.36) transverse process (Eberth and Currie, 2010)
(RTMP 98.63.37) pedal phalanx IV-1 (84 mm) (Eberth and Currie, 2010)
(RTMP 98.63.38) tooth (41 mm) (Eberth and Currie, 2010)
(RTMP 98.63.41) pedal phalanx II-1 (111 mm) (Eberth and Currie, 2010)
(RTMP 98.63.42) gastralia (Eberth and Currie, 2010)
(RTMP 98.63.44) premaxillary tooth (28 mm) (Eberth and Currie, 2010)
(RTMP 98.63.45) tooth (78 mm) (Eberth and Currie, 2010)
(RTMP 98.63.46) tooth (44 mm) (Eberth and Currie, 2010)
(RTMP 98.63.49) anterior dentary tooth (Eberth and Currie, 2010)
(RTMP 98.63.50) tooth (Eberth and Currie, 2010)
(RTMP 98.63.51) rib (Eberth and Currie, 2010)
(RTMP 98.63.52) tooth (44 mm) (Eberth and Currie, 2010)
(RTMP 98.63.53) tooth (49 mm) (Eberth and Currie, 2010)
(RTMP 98.63.54) tooth (Eberth and Currie, 2010)
(RTMP 98.63.55) tooth (26 mm) (Eberth and Currie, 2010)
(RTMP 98.63.56) rib (Eberth and Currie, 2010)
(RTMP 98.63.57) manual phalanx II-1 (68 mm) (Eberth and Currie, 2010)
(RTMP 98.63.58) ectopterygoid (Eberth and Currie, 2010)
(RTMP 98.63.59) tooth (Eberth and Currie, 2010)
(RTMP 98.63.60) ungual (Eberth and Currie, 2010)
(RTMP 98.63.61) limb element (Eberth and Currie, 2010)
(RTMP 98.63.65) tooth (Eberth and Currie, 2010)
(RTMP 98.63.67) tooth (65 mm) (Eberth and Currie, 2010)
(RTMP 98.63.68) vertebra (Eberth and Currie, 2010)
(RTMP 98.63.73) ischium (Eberth and Currie, 2010)
(RTMP 98.63.74) rib (Eberth and Currie, 2010)
(RTMP 98.63.75) phalanx (Eberth and Currie, 2010)
(RTMP 98.63.76) rib (670 mm) (Eberth and Currie, 2010)
(RTMP 98.63.77) splenial (Eberth and Currie, 2010)
(RTMP 98.63.78) tooth (Eberth and Currie, 2010)
(RTMP 98.63.81) rib (Eberth and Currie, 2010)
(RTMP 98.63.82) surangular, angular, articular (Eberth and Currie, 2010)
(RTMP 98.63.83) (juvenile) premaxilla (Eberth and Currie, 2010)
(RTMP 98.63.84) quadratojugal (Eberth and Currie, 2010)
(RTMP 98.63.85) surangular (Eberth and Currie, 2010)
(RTMP 98.63.87) vomer (Currie, 2003)
(RTMP 98.63.88) maxilla (Carr, 2010)
(RTMP 98.63.89) scapula (Eberth and Currie, 2010)
(RTMP 98.63.90) ischium (695 mm) (Eberth and Currie, 2010)
(RTMP 98.63.91) tooth (Eberth and Currie, 2010)
(RTMP 98.63.92) fragment (Eberth and Currie, 2010)
(RTMP 98.63.93) manual phalanx (Eberth and Currie, 2010)
(RTMP 98.63.94) tooth (Eberth and Currie, 2010)
(RTMP 98.63.95) gastralia (Eberth and Currie, 2010)
(RTMP 98.63.96) partial gastralium (Bell, 2010)
(RTMP 98.63.98) ungual (Eberth and Currie, 2010)
(RTMP 98.63.99) pedal phalanx I-1 (Eberth and Currie, 2010)
(RTMP 98.64.5) maxilla (Eberth and Currie, 2010)
(RTMP 98.64.6) phalanx (Eberth and Currie, 2010)
(RTMP 98.64.7) nasal (Eberth and Currie, 2010)
(RTMP 98.64.8) cranial element (Eberth and Currie, 2010)
(RTMP 98.64.9) quadrate (Eberth and Currie, 2010)
(RTMP 98.64.10) nasal (Eberth and Currie, 2010)
(RTMP 98.64.11) maxilla (Eberth and Currie, 2010)
(RTMP 98.64.12) cranial element (Eberth and Currie, 2010)
(RTMP 98.68.159) phalanx (Eberth and Currie, 2010)
(RTMP 99.50.1) tibia (Eberth and Currie, 2010)
(RTMP 99.50.2) (21 year old) fibula (Eberth and Currie, 2010)
(RTMP 99.50.3) femur (Eberth and Currie, 2010)
(RTMP 99.50.4) tibia (Eberth and Currie, 2010)
(RTMP 99.50.5) maxilla (Eberth and Currie, 2010)
(RTMP 99.50.6) element (Eberth and Currie, 2010)
(RTMP 99.50.7) vertebra (Eberth and Currie, 2010)
(RTMP 99.50.8) scapula (Eberth and Currie, 2010)
(RTMP 99.50.9) vertebra (Eberth and Currie, 2010)
(RTMP 99.50.10) maxilla (Eberth and Currie, 2010)
(RTMP 99.50.12) partial metatarsal III (Eberth and Currie, 2010)
(RTMP 99.50.13) vertebra (Eberth and Currie, 2010)
(RTMP 99.50.14) pedal phalanx II-1 (Bell, 2010)
(RTMP 99.50.15) phalanx (Eberth and Currie, 2010)
(RTMP 99.50.16) phalanx (Eberth and Currie, 2010)
(RTMP 99.50.17) palatine (Eberth and Currie, 2010)
(RTMP 99.50.18) caudal vertebra (Eberth and Currie, 2010)
(RTMP 99.50.19) (6 year old juvenile) femur (Eberth and Currie, 2010)
(RTMP 99.50.20) dentary (Eberth and Currie, 2010)
(RTMP 99.50.21) phalanx (Eberth and Currie, 2010)
(RTMP 99.50.22) parietal (Eberth and Currie, 2010)
(RTMP 99.50.24) metatarsal IV (460 mm) (Eberth and Currie, 2010)
(RTMP 99.50.25) metatarsal III (474 mm) (Eberth and Currie, 2010)
(RTMP 99.50.26) (13 year old) metatarsal II (434 mm) (Eberth and Currie, 2010)
(RTMP 99.50.27) astragalus (85 mm wide) (Eberth and Currie, 2010)
(RTMP 99.50.28) (12 year old) pedal phalanx II-2? (57 mm) (Eberth and Currie,
2010)
(RTMP 99.50.29) vertebra (Eberth and Currie, 2010)
(RTMP 99.50.30) gastralia (Eberth and Currie, 2010)
(RTMP 99.50.31) rib (Eberth and Currie, 2010)
(RTMP 99.50.32) metatarsal V (202 mm) (Eberth and Currie, 2010)
(RTMP 99.50.33) ungual (Eberth and Currie, 2010)
(RTMP 99.50.34) ungual (Eberth and Currie, 2010)
(RTMP 99.50.35) ungual (Eberth and Currie, 2010)
(RTMP 99.50.36) (juvenile) basioccipital (Eberth and Currie, 2010)
(RTMP 99.50.37) pedal phalanx III-2? (Eberth and Currie, 2010)
(RTMP 99.50.38) chevron (Eberth and Currie, 2010)
(RTMP 99.50.39) metacarpal (28 mm) (Eberth and Currie, 2010)
(RTMP 99.50.40) dentary (385 mm) (Eberth and Currie, 2010)
(RTMP 99.50.41) rib (365 mm) (Eberth and Currie, 2010)
(RTMP 99.50.42) rib (570 mm) (Eberth and Currie, 2010)
(RTMP 99.50.43) rib (Eberth and Currie, 2010)
(RTMP 99.50.44) pedal phalanx II-2 (82 mm) (Eberth and Currie, 2010)
(RTMP 99.50.45) gastralia (295 mm) (Eberth and Currie, 2010)
(RTMP 99.50.46) chevron (Eberth and Currie, 2010)
(RTMP 99.50.47) (juvenile) premaxilla (Eberth and Currie, 2010)
(RTMP 99.50.48) pedal phalanx II-1 (113 mm) (Eberth and Currie, 2010)
(RTMP 99.50.49) caudal vertebra (64 mm) (Eberth and Currie, 2010)
(RTMP 99.50.50) gastralia (Eberth and Currie, 2010)
(RTMP 99.50.51) rib (Eberth and Currie, 2010)
(RTMP 99.50.52) femur (Eberth and Currie, 2010)
(RTMP 99.50.53) pedal phalanx III-2 (81 mm) (Eberth and Currie, 2010)
(RTMP 99.50.54) element (Eberth and Currie, 2010)
(RTMP 99.50.55) pedal phalanx III-2 (Eberth and Currie, 2010)
(RTMP 99.50.56) cranial element (Eberth and Currie, 2010)
(RTMP 99.50.57) lacrimal (Carr, 2010)
(RTMP 99.50.58) premaxillary tooth (Eberth and Currie, 2010)
(RTMP 99.50.59) tooth (36 mm) (Eberth and Currie, 2010)
(RTMP 99.50.60) tooth (61 mm) (Eberth and Currie, 2010)
(RTMP 99.50.61) premaxillary tooth (Eberth and Currie, 2010)
(RTMP 99.50.62) premaxillary tooth (Eberth and Currie, 2010)
(RTMP 99.50.63) tooth (50 mm) (Eberth and Currie, 2010)
(RTMP 99.50.64) tooth (Eberth and Currie, 2010)
(RTMP 99.50.65) tooth (48 mm) (Eberth and Currie, 2010)
(RTMP 99.50.66) tooth (30 mm) (Eberth and Currie, 2010)
(RTMP 99.50.67) mid maxillary tooth (40 mm) (Reichel, 2010)
(RTMP 99.50.68) tooth (45 mm) (Eberth and Currie, 2010)
(RTMP 99.50.69) tooth (24 mm) (Eberth and Currie, 2010)
(RTMP 99.50.70) tooth (19 mm) (Eberth and Currie, 2010)
(RTMP 99.50.71) tooth (Eberth and Currie, 2010)
(RTMP 99.50.72) tooth (Eberth and Currie, 2010)
(RTMP 99.50.73) tooth (Eberth and Currie, 2010)
(RTMP 99.50.74) tooth (70 mm) (Eberth and Currie, 2010)
(RTMP 99.50.75) tooth (51 mm) (Eberth and Currie, 2010)
(RTMP 99.50.76) tooth (Eberth and Currie, 2010)
(RTMP 99.50.77) tooth (56 mm) (Eberth and Currie, 2010)
(RTMP 99.50.78) premaxillary tooth (Eberth and Currie, 2010)
(RTMP 99.50.79) (juvenile) tooth (Eberth and Currie, 2010)
(RTMP 99.50.80) tooth (48 mm) (Eberth and Currie, 2010)
(RTMP 99.50.81) tooth (63 mm) (Eberth and Currie, 2010)
(RTMP 99.50.82) tooth (Eberth and Currie, 2010)
(RTMP 99.50.83) tooth (25 mm) (Eberth and Currie, 2010)
(RTMP 99.50.84) tooth (67 mm) (Eberth and Currie, 2010)
(RTMP 99.50.85) tooth (55x21x? mm) (Eberth and Currie, 2010)
(RTMP 99.50.86) mid dentary tooth (47 mm) (Reichel, 2010)
(RTMP 99.50.87) tooth (Eberth and Currie, 2010)
(RTMP 99.50.88) tooth (Eberth and Currie, 2010)
(RTMP 99.50.89) (juvenile) maxillary tooth (Eberth and Currie, 2010)
(RTMP 99.50.90) tooth (Eberth and Currie, 2010)
(RTMP 99.50.91) tooth (62 mm) (Eberth and Currie, 2010)
(RTMP 99.50.92) tooth (60 mm) (Eberth and Currie, 2010)
(RTMP 99.50.93) tooth (Eberth and Currie, 2010)
(RTMP 99.50.94) tooth (42 mm) (Eberth and Currie, 2010)
(RTMP 99.50.95) dentary tooth (42 mm) (Eberth and Currie, 2010)
(RTMP 99.50.96) tooth (18 mm) (Eberth and Currie, 2010)
(RTMP 99.50.97) tooth (40 mm) (Eberth and Currie, 2010)
(RTMP 99.50.98) tooth (Eberth and Currie, 2010)
(RTMP 99.50.99) tooth (45 mm) (Eberth and Currie, 2010)
(RTMP 99.50.100) tooth (49 mm) (Eberth and Currie, 2010)
(RTMP 99.50.101) tooth (43 mm) (Eberth and Currie, 2010)
(RTMP 99.50.102) tooth (Eberth and Currie, 2010)
(RTMP 99.50.103) tooth (51 mm) (Eberth and Currie, 2010)
(RTMP 99.50.104) tooth (30 mm) (Eberth and Currie, 2010)
(RTMP 99.50.105) tooth (Eberth and Currie, 2010)
(RTMP 99.50.106) tooth (Eberth and Currie, 2010)
(RTMP 99.50.107) tooth (Eberth and Currie, 2010)
(RTMP 99.50.108) tooth (55 mm) (Eberth and Currie, 2010)
(RTMP 99.50.109) tooth (Eberth and Currie, 2010)
(RTMP 99.50.118) tooth (Eberth and Currie, 2010)
(RTMP 99.50.133) tooth (Eberth and Currie, 2010)
(RTMP 99.50.134) astragalus (Eberth and Currie, 2010)
(RTMP 99.50.135) elements (Eberth and Currie, 2010)
(RTMP 99.50.136) metacarpal (Eberth and Currie, 2010)
(RTMP 99.50.140) maxilla (Carr, 2010)
(RTMP 99.50.155) proximal rib (Eberth and Currie, 2010)
(RTMP 99.50.156) tooth (Eberth and Currie, 2010)
(RTMP 99.50.157) tooth (60 mm) (Eberth and Currie, 2010)
(RTMP 99.50.158) posterior dentary tooth (21 mm) (Reichel, 2010)
(RTMP 99.50.159) tooth (31 mm) (Eberth and Currie, 2010)
(RTMP 99.50.160) tooth (42 mm) (Eberth and Currie, 2010)
(RTMP 99.50.161) tooth (Eberth and Currie, 2010)
(RTMP 99.50.162) tooth (Eberth and Currie, 2010)
(RTMP 99.50.163) calcaneum (Eberth and Currie, 2010)
(RTMP 99.50.167) tooth (Eberth and Currie, 2010)
(RTMP 99.50.168) metacarpal (Eberth and Currie, 2010)
(RTMP 99.50.169) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.1) pedal ungual (Eberth and Currie, 2010)
(RTMP 2000.45.2) pedal ungual (Eberth and Currie, 2010)
(RTMP 2000.45.3) pedal ungual (Eberth and Currie, 2010)
(RTMP 2000.45.4) premaxillary tooth (24 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.5) (juvenile) tooth (19 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.6) tooth (28 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.7) (9 year old juvenile) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2000.45.9) (23 year old) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.10) tooth (46 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.11) pedal ungual (110 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.14) tooth (36 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.15) pedal phalanx IV-2 (72 mm) (Bell, 2010)
(RTMP 2000.45.16) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.17) tooth (52 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.18) tooth (41 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.19) pedal phalanx II-2? (75 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.20) angular (Eberth and Currie, 2010)
(RTMP 2000.45.21) pedal phalanx II-2? (106 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.22) gastralia (Eberth and Currie, 2010)
(RTMP 2000.45.23) tooth (23 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.25) (juvenile) premaxillary tooth (38 mm) (Eberth and Currie,
2010)
(RTMP 2000.45.26) lacrimal (Carr, 2010)
(RTMP 2000.45.27) spenial (Eberth and Currie, 2010)
(RTMP 2000.45.28) angular (Eberth and Currie, 2010)
(RTMP 2000.45.30) chevron (Eberth and Currie, 2010)
(RTMP 2000.45.32) element (Eberth and Currie, 2010)
(RTMP 2000.45.33) lateral gastralium (Eberth and Currie, 2010)
(RTMP 2000.45.34) cervical rib (Eberth and Currie, 2010)
(RTMP 2000.45.39) rib (Eberth and Currie, 2010)
(RTMP 2000.45.43) palatine (Carr, 2010; = RTMP 2000.45.93?) or tooth (Eberth
and Currie, 2010)
(RTMP 2000.45.44) pedal phalanx II-1 (151 mm) (Bell, 2010)
(RTMP 2000.45.45) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.46) pedal phalanx III-2 (Eberth and Currie, 2010)
(RTMP 2000.45.47) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.49) tooth (36 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.50) element (Eberth and Currie, 2010)
(RTMP 2000.45.53) tooth (65 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.54) tooth (22 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.55) tooth (52 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.56) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.58) phalanx (Eberth and Currie, 2010)
(RTMP 2000.45.59) tooth (35 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.60) rib (Eberth and Currie, 2010)
(RTMP 2000.45.62) coronoid-supradentary (Eberth and Currie, 2010)
(RTMP 2000.45.63) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.64) premaxillary tooth (Eberth and Currie, 2010)
(RTMP 2000.45.65) lacrimal (Carr, 2010)
(RTMP 2000.45.66) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.67) phalanx (Eberth and Currie, 2010)
(RTMP 2000.45.68) cervical rib (Eberth and Currie, 2010)
(RTMP 2000.45.69) distal caudal vertebra (Eberth and Currie, 2010)
(RTMP 2000.45.70) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.71) hyoid (Eberth and Currie, 2010)
(RTMP 2000.45.72) rib (Eberth and Currie, 2010)
(RTMP 2000.45.73) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.74) lacrimal (Carr, 2010; = RTMP 2001.45.74?) or rib (Eberth and
Currie, 2010)
(RTMP 2000.45.76) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.77) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.78) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.79) phalanx (Eberth and Currie, 2010)
(RTMP 2000.45.83) gastralia (Eberth and Currie, 2010)
(RTMP 2000.45.84) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.88) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.89) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.92) maxilla (Carr, 2010)
(RTMP 2000.45.93) palatine (Eberth and Currie, 2010)
(RTMP 2000.45.94) splenial (Eberth and Currie, 2010)
(RTMP 2000.45.95) dorsal vertebra (Eberth and Currie, 2010)
(RTMP 2000.45.96) astragalus (Eberth and Currie, 2010)
(RTMP 2000.45.99) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.100) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.106) cranial element (Eberth and Currie, 2010)
(RTMP 2000.45.107) gastralia (Eberth and Currie, 2010)
(RTMP 2000.45.108) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.109) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.110) tooth (Eberth and Currie, 2010)
(RTMP 2000.54.1) incomplete metatarsal I, incomplete phalanx I-1, pedal ungual
I, partial metatarsal II, incomplete phalanx II-1, phalanx II-2, partial pedal
ungual II, incomplete metatarsal III, partial phalanx III-1, phalanx III-2 or
3, incomplete pedal ungual III, partial metatarsal IV, two phalanges IV-?, four
pedal phalangeal fragments, two pedal ungual fragments, fragments (Tanke and
Currie, 2010)
(RTMP 2001.45.1) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.2) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.3) tooth (37 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.4) partial tooth (Eberth and Currie, 2010)
(RTMP 2001.45.5) tooth fragment (Eberth and Currie, 2010)
(RTMP 2001.45.6) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.7) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.8) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.9) partial tooth (Eberth and Currie, 2010)
(RTMP 2001.45.10) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.11) tooth (26 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.12) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.13) tooth (20 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.14) tooth (29 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.15) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.16) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.17) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.18) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.19) tooth (35 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.20) tooth (32 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.21) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.22) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.23) tooth (46 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.24) tooth (30 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.25) tooth (44 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.26) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.27) tooth (17 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.28) premaxillary tooth (35 mm) (Reichel, 2010)
(RTMP 2001.45.29) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.30) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.31) tooth (37 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.32) tooth (39 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.33) tooth (39 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.34) tooth (57 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.35) tooth (71 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.36) tooth (61 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.37) tooth (55 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.38) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.39) tooth (Eberth and Currie, 2010)
(RTMP 2001.45.40) tooth (24 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.41) tooth (22 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.42) pedal ungual (Eberth and Currie, 2010)
(RTMP 2001.45.43) ungual (Eberth and Currie, 2010)
(RTMP 2001.45.44) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2001.45.45) phalanx (Eberth and Currie, 2010)
(RTMP 2001.45.46) pedal phalanx II-2 (Eberth and Currie, 2010)
(RTMP 2001.45.47) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2001.45.48) pedal phalanx IV-2 (74 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.49) (16 year old) pedal phalanx II-1 (130 mm) (Eberth and Currie,
2010)
(RTMP 2001.45.50) phalanx (Eberth and Currie, 2010)
(RTMP 2001.45.51) astragalus (Eberth and Currie, 2010)
(RTMP 2001.45.52) metacarpal (Eberth and Currie, 2010)
(RTMP 2001.45.53) metatarsal (Eberth and Currie, 2010)
(RTMP 2001.45.54) metatarsal (Eberth and Currie, 2010)
(RTMP 2001.45.56) vertebra (Eberth and Currie, 2010)
(RTMP 2001.45.57) vertebra (Eberth and Currie, 2010)
(RTMP 2001.45.58) caudal vertebra (24 mm) (Eberth and Currie, 2010)
(RTMP 2001.45.59) neural arch (Eberth and Currie, 2010)
(RTMP 2001.45.60) (14 year old) metatarsal IV (Eberth and Currie, 2010)
(RTMP 2001.45.61) proximal dorsal rib (Eberth and Currie, 2010)
(RTMP 2001.45.62) rib (Eberth and Currie, 2010)
(RTMP 2001.45.63) rib (Eberth and Currie, 2010)
(RTMP 2001.45.64) rib (Eberth and Currie, 2010)
(RTMP 2001.45.65) rib (Eberth and Currie, 2010)
(RTMP 2001.45.66) rib (Eberth and Currie, 2010)
(RTMP 2001.45.67) rib (Eberth and Currie, 2010)
(RTMP 2001.45.68) gastralia (Eberth and Currie, 2010)
(RTMP 2001.45.69) gastralia (Eberth and Currie, 2010)
(RTMP 2001.45.70) axial rib (Eberth and Currie, 2010)
(RTMP 2001.45.71) furcula (Eberth and Currie, 2010)
(RTMP 2001.45.72) rib or scapulocoracoid (Eberth and Currie, 2010)
(RTMP 2001.45.73) scapula (Eberth and Currie, 2010)
(RTMP 2001.45.74) lacrimal (Eberth and Currie, 2010)
(RTMP 2001.45.75) quadrate (Eberth and Currie, 2010)
(RTMP 2001.45.77) surangular (Eberth and Currie, 2010)
(RTMP 2001.45.78) maxilla (Eberth and Currie, 2010)
(RTMP 2002.45.1) tooth (37 mm) (Eberth and Currie, 2010)
(RTMP 2002.45.2) tooth (26 mm) (Eberth and Currie, 2010)
(RTMP 2002.45.3) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.4) tooth fragment (Eberth and Currie, 2010)
(RTMP 2002.45.5) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.6) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.7) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.8) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.9) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.10) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.11) tooth (40 mm) (Eberth and Currie, 2010)
(RTMP 2002.45.12) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.13) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.14) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.15) tooth (58 mm) (Eberth and Currie, 2010)
(RTMP 2002.45.16) tooth (62 mm) (Eberth and Currie, 2010)
(RTMP 2002.45.17) tooth (26 mm) (Eberth and Currie, 2010)
(RTMP 2002.45.18) hyoid (Eberth and Currie, 2010)
(RTMP 2002.45.19) splenial (Eberth and Currie, 2010)
(RTMP 2002.45.20) ilium (Eberth and Currie, 2010)
(RTMP 2002.45.21) dentary (Eberth and Currie, 2010)
(RTMP 2002.45.22) cranial element (Eberth and Currie, 2010)
(RTMP 2002.45.23) vertebra (Eberth and Currie, 2010)
(RTMP 2002.45.24) cervical vertebra (Eberth and Currie, 2010)
(RTMP 2002.45.25) vertebra (Eberth and Currie, 2010)
(RTMP 2002.45.26) vertebra (Eberth and Currie, 2010)
(RTMP 2002.45.27) caudal vertebra (Eberth and Currie, 2010)
(RTMP 2002.45.29) neural arch (Eberth and Currie, 2010)
(RTMP 2002.45.30) vertebra (Eberth and Currie, 2010)
(RTMP 2002.45.31) gastralia (Eberth and Currie, 2010)
(RTMP 2002.45.32) lateral gastralium (Eberth and Currie, 2010)
(RTMP 2002.45.33) cervical rib (Eberth and Currie, 2010)
(RTMP 2002.45.34) rib (Eberth and Currie, 2010)
(RTMP 2002.45.35) cervical rib (Eberth and Currie, 2010)
(RTMP 2002.45.36) rib (Eberth and Currie, 2010)
(RTMP 2002.45.37) rib (Eberth and Currie, 2010)
(RTMP 2002.45.38) rib (Eberth and Currie, 2010)
(RTMP 2002.45.39) rib (Eberth and Currie, 2010)
(RTMP 2002.45.41) rib (Eberth and Currie, 2010)
(RTMP 2002.45.42) phalanx (Eberth and Currie, 2010)
(RTMP 2002.45.43) humerus (Eberth and Currie, 2010)
(RTMP 2002.45.44) phalanx (Eberth and Currie, 2010)
(RTMP 2002.45.45) ilium (Eberth and Currie, 2010)
(RTMP 2002.45.46; = RTMP 2002.5.46 in Erickson et al., 2010?) (50.3 kg; 2 year
old juvenile) fibula (Eberth and Currie, 2010)
(RTMP 2002.45.47) metatarsal (Eberth and Currie, 2010)
(RTMP 2002.45.63; = RTMP 99.50.2?) fibula (Eberth and Currie, 2010)
(RTMP 2002.45.64) metacarpal III (Eberth and Currie, 2010)
(RTMP 2002.45.65) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2002.45.66) vertebra (Eberth and Currie, 2010)
(RTMP 2002.45.67) tooth (Eberth and Currie, 2010)
(RTMP 2002.45.68) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.1) tooth (29 mm) (Eberth and Currie, 2010)
(RTMP 2003.45.2) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.3) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.4) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.5) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.6) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.7) tooth (75 mm) (Eberth and Currie, 2010)
(RTMP 2003.45.8) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.9) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.10) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.11) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.12) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.13) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.14) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.15) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.16) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.17) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.18) gastralia (Eberth and Currie, 2010)
(RTMP 2003.45.19) gastralia (Eberth and Currie, 2010)
(RTMP 2003.45.20) lateral gastralium (Eberth and Currie, 2010)
(RTMP 2003.45.21) mandibular element (Eberth and Currie, 2010)
(RTMP 2003.45.23) vertebra (Eberth and Currie, 2010)
(RTMP 2003.45.24) vertebra (Eberth and Currie, 2010)
(RTMP 2003.45.25) rib (Eberth and Currie, 2010)
(RTMP 2003.45.26) scapula (Eberth and Currie, 2010)
(RTMP 2003.45.27) rib (Eberth and Currie, 2010)
(RTMP 2003.45.28) cervical rib (Eberth and Currie, 2010)
(RTMP 2003.45.29) proximal rib (Eberth and Currie, 2010)
(RTMP 2003.45.30) rib (Eberth and Currie, 2010)
(RTMP 2003.45.31) rib (Eberth and Currie, 2010)
(RTMP 2003.45.32) pedal ungual (Eberth and Currie, 2010)
(RTMP 2003.45.33) manual ungual (Eberth and Currie, 2010)
(RTMP 2003.45.34) metatarsal (250 mm), phalanx (55 mm), phalanx (62 mm) (Eberth
and Currie, 2010)
(RTMP 2003.45.35) pedal phalanx III-2 (68 mm) (Eberth and Currie, 2010)
(RTMP 2003.45.36) metatarsal III (Eberth and Currie, 2010)
(RTMP 2003.45.37) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.38) tooth (Eberth and Currie, 2010)
(RTMP 2003.45.73) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2003.45.74) rib (Eberth and Currie, 2010)
(RTMP 2003.45.76) dentary (Eberth and Currie, 2010)
(RTMP 2003.45.79) rib (Eberth and Currie, 2010)
(RTMP 2003.45.81) rib (Eberth and Currie, 2010)
(RTMP 2003.45.83) lacrimal (Eberth and Currie, 2010)
(RTMP 2003.45.84) dentary (Bell, 2010)
(RTMP 2003.45.85) femur (Eberth and Currie, 2010)
(RTMP 2003.45.86) lateral gastralium (Eberth and Currie, 2010)
(RTMP 2003.45.87) metatarsal (Eberth and Currie, 2010)
(RTMP 2003.45.88) skin impression (Currie and Koppelhus, 2010)
(RTMP 2004.45.15) (4 year old) (Erickson et al., 2010)
(RTMP 2004.56.1) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.2) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.3) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.4) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.5) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.6) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.7) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.8) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.9) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.10) (juvenile) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.11) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.12) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.13) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.14) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.15) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.16) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.17) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.18) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.19) (juvenile) posterior maxillary tooth (Reichel, 2010)
(RTMP 2004.56.20) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.21) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.22) premaxillary tooth (Eberth and Currie, 2010)
(RTMP 2004.56.23) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.24) tooth (Eberth and Currie, 2010)
(RTMP 2004.56.25) nasal (Eberth and Currie, 2010)
(RTMP 2004.56.26) manual ungual (Eberth and Currie, 2010)
(RTMP 2004.56.27) radius (Eberth and Currie, 2010)
(RTMP 2004.56.28) gastralia (Eberth and Currie, 2010)
(RTMP 2004.56.29) gastralia (Eberth and Currie, 2010)
(RTMP 2004.56.30) cervical rib (Eberth and Currie, 2010)
(RTMP 2004.56.31) rib (Eberth and Currie, 2010)
(RTMP 2004.56.32) rib (Eberth and Currie, 2010)
(RTMP 2004.56.33) rib (Eberth and Currie, 2010)
(RTMP 2004.56.34) proximal rib (Eberth and Currie, 2010)
(RTMP 2004.56.35) rib (Eberth and Currie, 2010)
(RTMP 2004.56.36) rib (Eberth and Currie, 2010)
(RTMP 2004.56.37) rib (Eberth and Currie, 2010)
(RTMP 2004.56.38) rib (Eberth and Currie, 2010)
(RTMP 2004.56.39) rib (Eberth and Currie, 2010)
(RTMP 2004.56.40) vertebra (Eberth and Currie, 2010)
(RTMP 2004.56.41) chevron (Eberth and Currie, 2010)
(RTMP 2004.56.42) fibula (680 mm) (Eberth and Currie, 2010)
(RTMP 2004.56.43) (14 year old) metatarsal III (Eberth and Currie, 2010)
(RTMP 2004.56.44) metatarsal II or IV (Eberth and Currie, 2010)
(RTMP 2004.56.45) metatarsal III (Eberth and Currie, 2010)
(RTMP 2004.56.46) metatarsal II (Eberth and Currie, 2010)
(RTMP 2004.56.47) metatarsal (350 mm) (Eberth and Currie, 2010)
(RTMP 2004.56.48) (28 year old) pedal phalanx II-2 (120 mm) (Eberth and Currie,
2010)
(RTMP 2004.56.49) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2004.56.50) pedal phalanx I-1 (Eberth and Currie, 2010)
(RTMP 2004.56.51) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2004.56.52) pedal phalanx II-2 (Eberth and Currie, 2010)
(RTMP 2004.56.53) pedal phalanx (Eberth and Currie, 2010)
(RTMP 2004.56.54) pedal phalanx (150 mm) (Eberth and Currie, 2010)
(RTMP 2004.56.55) pedal phalanx (135 mm) (Eberth and Currie, 2010)
(RTMP 2004.56.56) pedal phalanx III-3 (76 mm) (Eberth and Currie, 2010)
(RTMP 2004.56.57) pedal phalanx III-1, phalanx III-2 (Eberth and Currie, 2010)
(RTMP 2004.56.67) pubis (Eberth and Currie, 2010)
(RTMP 2004.56.68) rib (891 mm) (Eberth and Currie, 2010)
(RTMP 2004.56.70) metatarsal (Eberth and Currie, 2010)
(RTMP 2005.50.1) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.2) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.3) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.4) tooth (60 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.5) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.6) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.7) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.8) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.9) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.10) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.11) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.13) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.14) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.15) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.16) tooth (37 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.17) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.18) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.19) tooth (22 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.20) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.21) angular (Eberth and Currie, 2010)
(RTMP 2005.50.22) surangular (Eberth and Currie, 2010)
(RTMP 2005.50.23) cranial element (Eberth and Currie, 2010)
(RTMP 2005.50.24) nasal (Eberth and Currie, 2010)
(RTMP 2005.50.25) maxilla (Eberth and Currie, 2010)
(RTMP 2005.50.26) jugal (Eberth and Currie, 2010)
(RTMP 2005.50.27) epipterygoid (Eberth and Currie, 2010)
(RTMP 2005.50.28) quadrate (Eberth and Currie, 2010)
(RTMP 2005.50.29) barincase (Eberth and Currie, 2010)
(RTMP 2005.50.30) caudal vertebra (44 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.31) centrum (Eberth and Currie, 2010)
(RTMP 2005.50.32) cervical zygapophysis (Eberth and Currie, 2010)
(RTMP 2005.50.33) metatarsal (Eberth and Currie, 2010)
(RTMP 2005.50.34) metatarsal (Eberth and Currie, 2010)
(RTMP 2005.50.35) metatarsal III (Eberth and Currie, 2010)
(RTMP 2005.50.36) distal pedal phalanx (Eberth and Currie, 2010)
(RTMP 2005.50.37) pedal phalanx II-1 (90 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.38) pedal ungual (73 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.39) pedal phalanx II-1 (106 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.40) phalanx (Eberth and Currie, 2010)
(RTMP 2005.50.41) femur (900 mm) (Eberth and Currie, 2010)
(RTMP 2005.50.42) fibula (Eberth and Currie, 2010)
(RTMP 2005.50.43) furcula (Eberth and Currie, 2010)
(RTMP 2005.50.44) cervical rib (Eberth and Currie, 2010)
(RTMP 2005.50.45) rib (Eberth and Currie, 2010)
(RTMP 2005.50.46) rib (Eberth and Currie, 2010)
(RTMP 2005.50.47) rib (Eberth and Currie, 2010)
(RTMP 2005.50.48) proximal rib (Eberth and Currie, 2010)
(RTMP 2005.50.49) rib (Eberth and Currie, 2010)
(RTMP 2005.50.50) rib (Eberth and Currie, 2010)
(RTMP 2005.50.51) rib (Eberth and Currie, 2010)
(RTMP 2005.50.52) rib (Eberth and Currie, 2010)
(RTMP 2005.50.53) rib (Eberth and Currie, 2010)
(RTMP 2005.50.54) rib (Eberth and Currie, 2010)
(RTMP 2005.50.55) rib (Eberth and Currie, 2010)
(RTMP 2005.50.56) rib (Eberth and Currie, 2010)
(RTMP 2005.50.57) rib (Eberth and Currie, 2010)
(RTMP 2005.50.58) rib (Eberth and Currie, 2010)
(RTMP 2005.50.67) caudal centrum (Eberth and Currie, 2010)
(RTMP 2005.50.69) tooth (Eberth and Currie, 2010)
(RTMP 2005.50.70) rib (Eberth and Currie, 2010)
(RTMP 2005.50.71) vertebra (Eberth and Currie, 2010)
(RTMP 2005.50.72) element (Eberth and Currie, 2010)
(RTMP coll.) twenty-five fragmentary teeth (Ryan, Currie, Gardner, Vickaryous
and Lavigne, 2000)
(RTMP coll.; Dry Island bonebed) (unassociated) quadrate, tooth (Eberth and
Currie, 2010)
(RTMP or UALVP coll.; Dry Island bonebed) (unassociated) two premaxillary teeth,
maxilla, two maxillary fragments, squamosal (collected?), two quadrates (one
uncollected), 1-2 ectopterygoids, 2 pterygoids (1 uncollected?), paroccipital
process, splenial (298 mm), coronoid (uncollected), surangular, 5 cranial elements,
71 teeth (19-22 uncollected; 18, 28, 30, 32, 33, 40, 40, 42, 45, 45, 54, 60,
61, 77 mm), 2 cervical vertebrae (1 uncollected?), cervical vertebra, cervical
rib, dorsal vertebra (140 mm), 2 dorsal rib (1 uncollected), gastralia (some
uncollected), sacral vertebra, mid caudal vertebra (123 mm), two distal caudal
vertebrae (43.6, 51 mm), 5 caudal vertebrae (1 uncollected), caudal centrum,
14 vertebrae (1-3 uncollected), 7 chevrons (0-2 uncollected; 380, 78 mm), 6
centra (4-5 uncollected), 13 neural arches (9-11 uncollected), 34 ribs (11-12
uncollected; 76 mm), 19 proximal ribs (4-5 uncollected), 32 rib fragments (25-26
uncollected), coracoid, furcula, 2 radii (1 uncollected; 140 mm), 3 ulnae (116.5
mm), phalanx II-2 (55 mm), manual phalanx, 3 ilia (2 uncollected), 4 pubes (3
uncollected), 2 ischia (1 incomplete), 6 femora (730, 860 mm), tibiae (1 uncollected;
655 mm), tibial fragment (uncollected), distal tibia (collected?), 6 fibulae
(one incomplete; one proximal; 650 mm), 3 astragali (one partial; one uncollected),
2 phalanges I-1 (75.5 mm), phalanx II-1 (160 mm), 2 phalanges II-2 (96, 75 mm),
2 metatarsals III (uncollected), phalanx III-1 (uncollected; 133 mm), 3 phalanges
III-2 (95, 108 mm), 2 phalanges III-3 (one distal), phalanx IV-1, phalanx IV-2,
phalanx IV-4 (48 mm), 3 metatarsals (2 incomplete; 2 uncollected), 2 pedal phalanges
(37 mm; 1 uncollected), 5 pedal unguals, limb element, limb fragment (uncollected),
phalanx, 22 unidentified elements (2 uncollected) (Eberth and Currie, 2010)
(UAVLP 47899) tooth (Torices et al., 2014)
(UALVP 47900) tooth (Torices et al., 2014)
(UALVP 47901) tooth (Torices et al., 2014)
(UALVP 47902) tooth (Torices et al., 2014)
(UALVP 47903) tooth (54x22x13.8 mm) (Torices et al., 2014)
(UALVP 47904) tooth (Torices et al., 2014)
(UALVP 47905) tooth (Torices et al., 2014)
(UALVP 47906) tooth (40.3x18.7x13.7 mm) (Torices et al., 2014)
(UALVP 47908) tooth (59.5x30.1x20 mm) (Torices et al., 2014)
(UALVP 47934) tooth (?x32.4x26.4 mm) (Torices et al., 2014)
(UALVP 47936) tooth (46.6x27.9x18.3 mm) (Torices et al., 2014)
(UALVP 47937) tooth (28.5x15.6x11 mm) (Torices et al., 2014)
(UALVP 48922) tooth (39x23.7x13.1 mm) (Torices et al., 2014)
(UALVP 48923) tooth (34.8x16.5x12.8 mm) (Torices et al., 2014)
(UALVP 48924) tooth (?x33.6x17.2 mm) (Torices et al., 2014)
(UALVP 48926) tooth (59x23.8x13.9 mm) (Torices et al., 2014)
(UALVP 48944) tooth (Torices et al., 2014)
(UALVP 48966) tooth (Torices et al., 2014)
(UALVP 48994) tooth (?x2.6x? mm) (Torices et al., 2014)
(UALVP 50689) tooth (40.9x?x? mm) (Torices et al., 2014)
(UALVP 50691) tooth (63.4x31.2x18.9 mm) (Torices et al., 2014)
(UALVP 50692) tooth (46.6x21.8x15 mm) (Torices et al., 2014)
(UALVP 50693) tooth (?x16.2x9.2 mm) (Torices et al., 2014)
(UALVP 50694) tooth (22.2x14x9 mm) (Torices et al., 2014)
(UALVP 50695) tooth (61.9x30.3x17.2 mm) (Torices et al., 2014)
(UALVP 50696) tooth (?x13.1x7.8 mm) (Torices et al., 2014)
(UALVP 50697) tooth (37.4x19.1x14.5 mm) (Torices et al., 2014)
(UALVP 50698) tooth (56.9x29.4x19.4 mm) (Torices et al., 2014)
(UALVP 50701) tooth (57.2x24.4x18.9 mm) (Torices et al., 2014)
(UALVP 50737) tooth (39.3x?x12 mm) (Torices et al., 2014)
(UALVP 51957) tooth (?x?x18.9 mm) (Torices et al., 2014)
(UALVP 52016) pedal phalanx II-1 (79 mm) (Eberth and Currie, 2010)
(UALVP 52017) tooth (Eberth and Currie, 2010)
(UALVP 52018) tooth fragment (Eberth and Currie, 2010)
(UALVP 52019) postorbital (Eberth and Currie, 2010)
(UALVP 52020) pedal ungual I (60 mm) (Eberth and Currie, 2010)
(UALVP 52021) tooth (Eberth and Currie, 2010)
(UALVP 52022) jugal (Eberth and Currie, 2010)
(UALVP 52023) ulna (Eberth and Currie, 2010)
(UALVP 52024) proximal rib (Eberth and Currie, 2010)
(UALVP 52025) proximal rib (Eberth and Currie, 2010)
(UALVP 52026) pedal ungual (87 mm) (Eberth and Currie, 2010)
(UALVP 52027) pedal ungual (Eberth and Currie, 2010)
(UALVP 52028) tooth (Eberth and Currie, 2010)
(UALVP 52029) tooth (Eberth and Currie, 2010)
(UALVP 52030) jugal (Eberth and Currie, 2010)
(UALVP 52031) pedal phalanx III-3 (Eberth and Currie, 2010)
(UALVP 52033) coracoid (Eberth and Currie, 2010)
(UALVP 52034) pedal phalanx IV-1 (88 mm) (Eberth and Currie, 2010)
(UALVP 52035) metatarsal II ((Eberth and Currie, 2010)
(UALVP 52036) femora (one proximal) (Eberth and Currie, 2010)
(UALVP 52037) tibia (685 mm) (Eberth and Currie, 2010)
(UALVP 52038) tibia, fibulae (Eberth and Currie, 2010)
(UALVP 52039) (~5.1 m, 10 year old) proximal fibula (Erickson et al., 2010)
(UALVP 52040; = UALVP 52062?) centrum (Eberth and Currie, 2010)
(UALVP 52041) humerus (Eberth and Currie, 2010)
(UALVP 52042) tooth (Eberth and Currie, 2010)
(UALVP 52043) tooth (Eberth and Currie, 2010)
(UALVP 52044) prearticular (Eberth and Currie, 2010)
(UALVP 52045) pedal phalanx IV-4 (35 mm) (Eberth and Currie, 2010)
(UALVP 52046) caudal vertebra (Eberth and Currie, 2010)
(UALVP 52047) centrum (Eberth and Currie, 2010)
(UALVP 52048) quadratojugal (Eberth and Currie, 2010)
(UALVP 52049) metatarsal III (Eberth and Currie, 2010)
(UALVP 52050) caudal vertebra (Eberth and Currie, 2010)
(UALVP 52051) caudal vertebra (Eberth and Currie, 2010)
(UALVP 52052) neural arch (Eberth and Currie, 2010)
(UALVP 52053) ischium (Eberth and Currie, 2010)
(UALVP 52054) quadrate? (Eberth and Currie, 2010)
(UALVP 52055) ?cranial element (Eberth and Currie, 2010)
(UALVP 52056) pedal phalanx III-1 (Eberth and Currie, 2010)
(UALVP 52057) braincase (Eberth and Currie, 2010)
(UALVP 52058; = UALVP 52060?) maxillary fragment (Eberth and Currie, 2010)
(UALVP 52059) humerus (Eberth and Currie, 2010)
(UALVP 52060) (Eberth and Currie, 2010)
(UALVP 52061) pedal ungual (93 mm) (Eberth and Currie, 2010)
(UALVP 52062; = UALVP 52040?) centrum (Eberth and Currie, 2010)
(UALVP 52063) incomplete pubis (Eberth and Currie, 2010)
(UALVP 52064) sacrum (Eberth and Currie, 2010)
(UALVP 52065) distal fibula (Eberth and Currie, 2010)
(UALVP 52066) ulna (Eberth and Currie, 2010)
(UALVP 52067) ulna (Eberth and Currie, 2010)
(UALVP 52068) pubis? (Eberth and Currie, 2010)
(UALVP 52069) ectopterygoid (Eberth and Currie, 2010)
(UALVP 52070) pedal phalanx III-2 (78.4 mm) (Eberth and Currie, 2010)
(UALVP 52071) tooth (Eberth and Currie, 2010)
(UALVP 52072) cranial fragment (Eberth and Currie, 2010)
(UALVP 52073) centrum (Eberth and Currie, 2010)
(UALVP 52074) metatarsal IV (Eberth and Currie, 2010)
(UALVP 52077) gastralia (Eberth and Currie, 2010)
(UALVP 52078) (Eberth and Currie, 2010)
(UALVP 52079) ectopterygoid (Eberth and Currie, 2010)
(UALVP 52080) metatarsal (Eberth and Currie, 2010)
(UALVP 52081) metatarsal (Eberth and Currie, 2010)
(UALVP 52082) caudal vertebra (Eberth and Currie, 2010)
(UALVP 52083) cranial fragment (Eberth and Currie, 2010)
(UALVP 52084) metatarsal (Eberth and Currie, 2010)
(UALVP 52085) pedal ungual (107 mm) (Eberth and Currie, 2010)
(UALVP 52086) centrum (Eberth and Currie, 2010)
(UALVP 52087) surangular (Eberth and Currie, 2010)
(UALVP 52088) surangular (Eberth and Currie, 2010)
(UALVP 52090) pedal phalanx (Eberth and Currie, 2010)
(UALVP 52096) cranial element (Eberth and Currie, 2010)
(UALVP 52097) (Eberth and Currie, 2010)
(UALVP 52098) limb element (Eberth and Currie, 2010)
(UALVP 52099) (Eberth and Currie, 2010)
(UALVP 52101) (~8.1 m, 22 year old) distal tibia (width 230 mm) (Erickson et
al., 2010)
(UALVP 52102) pedal phalanx III-3 (Eberth and Currie, 2010)
(UALVP 52104) caudal vertebra (Eberth and Currie, 2010)
(UALVP 52106) rib (Eberth and Currie, 2010)
(UALVP 52107) tooth (Eberth and Currie, 2010)
(UALVP 52108) tooth (Eberth and Currie, 2010)
(UALVP 52110) tooth fragment (Eberth and Currie, 2010)
(UALVP 52111) caudal vertebra (Eberth and Currie, 2010)
(UALVP 52112) distal caudal vertebra (Eberth and Currie, 2010)
(UALVP 52115) proximal fibula (Eberth and Currie, 2010)
(UALVP 52117) pedal phalanx (Eberth and Currie, 2010)
(UALVP 52119) phalanx (Eberth and Currie, 2010)
(UALVP 52121) phalanx (Eberth and Currie, 2010)
(UALVP 52122) fibula (Eberth and Currie, 2010)
(UALVP 52123) manual phalanx (Eberth and Currie, 2010)
(UALVP 52124) manual ungual (Eberth and Currie, 2010)
(UALVP 52127) tooth (Eberth and Currie, 2010)
(UALVP 52128) ilium (860 mm) (Eberth and Currie, 2010)
(UALVP 52129) (~6.6 m, 16 year old) pubes (Erickson et al., 2010)
(UALVP 52130) cranial element, phalanx (Eberth and Currie, 2010)
(UALVP 52134) (~9.1 m, 26 year old) pedal phalanx IV-4 (Erickson et al., 2010)
(UALVP 52331) tooth (65.8x28.4x21.1 mm) (Torices et al., 2014)
(UALVP 52332) tooth (48.8x20.1x16.2 mm) (Torices et al., 2014)
(UALVP 52333) tooth (Torices et al., 2014)
(UALVP 52620) tooth (66.7x30.2x19.1 mm) (Torices et al., 2014)
(UALVP 52682) tooth (Torices et al., 2014)
(UALVP 53088) tooth (25.4x11.6x8.4 mm) (Torices et al., 2014)
(UALVP 53135) tooth (Torices et al., 2014)
(UALVP 53137) tooth (Torices et al., 2014)
(UALVP 53139) tooth (Torices et al., 2014)
(UALVP 53227) tooth (Torices et al., 2014)
(UALVP 53238) tooth (38.2x17x11.3 mm) (Torices et al., 2014)
(UALVP 53247) tooth 28.1x?x8.7 mm) (Torices et al., 2014)
(UALVP 53264) tooth (18.7x12.9x8.8 mm) (Torices et al., 2014)
(UALVP 53506) tooth (?x31.2x19.8 mm) (Torices et al., 2014)
(UALVP 54210) tooth (Torices et al., 2014)
(UALVP 54212) tooth (Torices et al., 2014)
(UALVP 54213) tooth (?x21.7x12 mm) (Torices et al., 2014)
(UALVP 54316) tooth (?x20.4x13.6 mm) (Torices et al., 2014)
(UALVP 54704) tooth (48.1x24.6x20.6 mm) (Torices et al., 2014)
(UALVP 54706) tooth (Torices et al., 2014)
(UALVP 54707) tooth (Torices et al., 2014)
(UALVP 54708) tooth (Torices et al., 2014)
(UALVP 54737) tooth (48.4x26.6x17.1 mm) (Torices et al., 2014)
(UALVP 54743) tooth (45.6x24.5x13.6 mm) (Torices et al., 2014)
(UALVP 54833) tooth (42.9x22.8x13.1 mm) (Torices et al., 2014)
(UALVP 54835) tooth (Torices et al., 2014)
(UALVP 54836) tooth (Torices et al., 2014)
(UALVP 54837) tooth (Torices et al., 2014)
(UALVP 54857) tooth (21.1x14.6x9 mm) (Torices et al., 2014)
(UALVP 58916) tooth (Torices et al., 2014)
(UALVP 59599) (~1 m; embryo) pedal ungual (10.0 mm) (Funston, Powers, Whitebone, Brusatte, Scannella, Horner and Currie, 2020)
(UALVP coll.) (embryo) premaxillary tooth (16 mm), teeth (Funston,
Powers, Whitebone, Brusatte, Scannella, Horner and Currie, 2020)
dentary, teeth (Ryan, Bell and Eberth, 1995)
material (Evans et al., 2003)
Diagnosis- (after Carpenter, 1992) interfenestral strut narrow.
(after Carr, 2010) medial pneumatic recess in lacrimal internal angle (also
in tyrannosaurines); posterior palatine pneumatic recess large and deeper than
anterior pneumatic recess; dorsoventrally deep anteromedial palatine process.
Comments- The holotype was discovered in 1884, and the paratype in 1889.
The these were initially referred to Laelaps incrassatus by Cope (1892),
which is based on two teeth from the earlier Judith River Group of Montana.
Lambe (1903, 1904) then published more detailed descriptions of these specimens
as Dryptosaurus incrassatus. The combination was first used by Hay (1902)
because Laelaps was found to be preoccupied and replaced with Dryptosaurus
by Marsh. However, Lambe's (1904) statement that Cope's original teeth and dentary
are more likely Deinodon, making the Horseshoe Canyon specimens the types
of incrassatus is incorrect. The name must stick with Cope's original
Judith River holotype. Osborn (1905) recognized this and created the taxon Albertosaurus
sarcophagus for the Horseshoe Canyon specimens, to distinguish them from
the Montanan type material of incrassatus. The holotype of Albertosaurus
arctunguis (ROM 807) was discovered in 1923 and described by Parks (1928),
but has since been synonymized with A. sarcophagus (Russell, 1970).
The Dry Island bonebed was discovered in 1910 and its initial collections are
catalogued as AMNH 5218 and 5226-5235 (Currie, 2000). Exactly which elements
are catalogued as AMNH 5218 differs between references- Currie lists the non-hindlimb
material entered above plus "two femora, three tibiae, half a fibula, two
astragali, one calcaneum, a pair of associated metatarsals (II-III), six isolated
metatarsals, 42 phalanges, and seven unguals", while Eberth and Currie
(2010) list two humeri, three femora, four tibiae, a fibula, two astragali,
a metatarsal I, 37 phalanges and one ungual. At least 26 individuals are represented
(Erickson et al., 2010), and RTMP (98.63, 98.64, 99.50, 2000.45, 2001.45, 2002.45,
2003.45, 2004.56, 2005.50 specimens) and UA (2006-2010; Eberth and Currie, 2010
specimens above) excavation has produced over a thousand elements, most of which
have yet to be described (Currie and Koppelhus, 2010; Eberth and Currie, 2010).
While Currie thought a large pedal phalanx III-3 catalogued in AMNH 5218 might
be Daspletosaurus, no other evidence of that genus is present making
this highly unlikely. Bell et al. (2017) described the skin impressions of RTMP
94.186.1.
Osborn (1916) referred AMNH 5255 questionably to Ornithomimus velox,
but this hindlimb is now identified as Tyrannosauridae on the AMNH online catalogue.
It may be Albertosaurus based on provenance, and is listed as Tetanurae indet. by Carrano (1998). CMN 11315 was first identified
as Daspletosaurus (Russell, 1970), but is Albertosaurus (Currie,
2003). Bell (2007) first identified the Danek bonebed of the Hoseshoe Canyon
Formation as including Daspletosaurus, though the material was later
found to be Albertosaurus (Bell and Currie, 2014; Torices et al., 2014).
Ford (paleofile.com) lists an RTMP specimen noted by Rondeau (1995) in a newspaper
article under Albertosaurus, but based on its locality information and
Tanke (DML, 1996) this would seem to be RTMP 91.163.1, which is a Gorgosaurus
specimen according to Currie (2003).
Funston et al. (2020) mention "a possible premaxilla ... from the
Horseshoe Canyon Formation of Alberta" as a possible embryonic
tyrannosaurid (presumably Albertosaurus
based on providence), but this specimen (UALVP coll.) turned out to be
more likely a fragment of embryonic troodontid dentary (Funston pers.
comm.).
References- Cope, 1892. Skull of the dinosaurian Laelaps incrassatus
Cope. Proceedings of the American Philosophical Society. 30, 240-245.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 1-868.
Lambe, 1903. The lower jaw of Dryptosaurus incrassatus (Cope). The Ottawa
Naturalist. 175, 133-139.
Lambe, 1904. On Dryptosaurus incrassatus (Cope), from the Edmonton Series
of the North West Territory. Geological Survey of Canada Contributions to Canadian
Palaeontology. 3(3), 1-27.
Osborn, 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs.
Bulletin of the American Museum of Natural History. 21, 259-265.
Lambe, 1914. Report of the vertebrate paleontologist. Summary report of the
Geological Survey, 1912. 396-403.
Sternberg, 1915. Field notes (copy at RTMP).
Osborn, 1916. Skeletal adaptation of Ornitholestes, Struthiomimus,
Tyrannosaurus. Bulletin of the American Museum of Natural History. 35,
733-771.
Russell and Chamney, 1967. Notes on the biostratigraphy of Dinosaurian and microfossil
faunas in the Edmonton Formation (Cretaceous), Alberta. National Museum of Canada
Natural History Papers. 35, 1-22.
Russell, 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National
Museum of Natural Science Publications in Palaeontology. 1, 1-34.
Coy, 1982. Field notes (copy at RTMP).
Maier, 1985. Field notes (copy at RTMP).
Danis, 1986. Field notes (copy at RTMP).
Carpenter, 1992. Tyrannosaurids (Dinosauria) of Asia and North America. In Mateer
and Chen (eds.). Aspects of nonmarine Cretaceous geology. Ocean Press. 250-268.
Erickson, 1995. Split carinae on tyrannosaurid teeth and implications of their
development. Journal of Vertebrate Paleontology. 15(2), 268-274.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku
Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.
Ryan, Bell and Eberth, 1995. Taphonomy of a hadrosaur (Ornithischia:Hadrosauridae)
bone bed from the Horseshoe Canyon Formation (Early Maastrichtian), Alberta,
Canada. Journal of Vertebrate Paleontology. 15(3), 51A.
Rondeau, 1995. Brownfield dinosaur finds new home. The Rebiem. 3-7-1995.
Tanke, DML 1996. https://web.archive.org/web/20191009080800/http://dml.cmnh.org/1996Apr/msg00243.html
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD thesis, The University of Chicago. 424
pp.
Makovicky and Currie, 1998. The presence of a furcula in tyrannosaurid theropods,
and its phylogenetic and functional implications. Journal of Vertebrate Paleontology.
18(1), 143-149.
Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria).
Journal of Vertebrate Paleontology.19(3), 497-520.
Currie, 2000. Possible evidence of gregarious behavior in tyrannosaurids. Gaia.
15, 271-277.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous
of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Evans, Lam, Maddin and Conacher, 2003. Taphonomy of the Prehistoric Park quarry,
Horseshoe Canyon Formation, Drumheller, Alberta. Alberta Palaeontological Society
Seventh Annual Symposium, Abstracts Volume. 25-28.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Bell, 2007. The Danek bonebed: An unusual dinosaur assemblage from the Horseshoe
Canyon Formation, Edmonton, Alberta. Journal of Vertebrate Paleontology. 27(3),
46A.
Wolff, Salisbury, Horner, Varricchio and Hansen, 2009. Common avian infection
plagued the tyrant dinosaurs. PLoS ONE. 4(9), e7288.
Bell, 2010. Palaeopathological changes in a population of Albertosaurus sarcophagus
from the Upper Cretaceous Horseshoe Canyon Formation of Alberta, Canada. Canadian
Journal of Earth Sciences. 47(9), 1263-1268.
Buckley, Larson, Reichel and Samman, 2010. Quantifying tooth variation within
a single population of Albertosaurus sarcophagus (Theropoda: Tyrannosauridae)
and implications for identifying isolated teeth of tyrannosaurids. Canadian
Journal of Earth Sciences. 47(9), 1227-1251.
Carr, 2010. A taxonomic assessment of the type series of Albertosaurus sarcophagus
and the identity of Tyrannosauridae (Dinosauria, Coelurosauria) in the Albertosaurus
bonebed from the Horseshoe Canyon Formation (Campanian-Maastrichtian, Late Cretaceous).
Canadian Journal of Earth Sciences. 47(9), 1213-1226.
Currie and Eberth, 2010. On gregarious behavior in Albertosaurus. Canadian
Journal of Earth Sciences. 47(9), 1277-1289.
Currie and Koppelhus, 2010. Introduction to Albertosaurus special issue.
Canadian Journal of Earth Sciences. 47(9), 1111-1114.
Eberth and Currie, 2010. Stratigraphy, sedimentology, and taphonomy of the Albertosaurus
bonebed (upper Horseshoe Canyon Formation; Maastrichtian), southern Alberta,
Canada. Canadian Journal of Earth Sciences. 47(9), 1119-1143.
Erickson, Currie, Inouye and Winn, 2010. A revised life table and survivorship
curve for Albertosaurus sarcophagus based on the Dry Island mass death
assemblage. Canadian Journal of Earth Sciences. 47(9), 1269-1275.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe
Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta,
with special reference to the Albertosaurus sarcophagus bonebed. Canadian
Journal of Earth Sciences. 47(9), 1159-1181.
Reichel, 2010. The heterodonty of Albertosaurus sarcophagus and Tyrannosaurus
rex: Biomechanical implications inferred through 3-D models. Canadian Journal
of Earth Sciences. 47(9), 1227-1251.
Tanke and Currie, 2010. A history of Albertosaurus discoveries in Alberta,
Canada. Canadian Journal of Earth Sciences. 47(9), 1197-1211.
Sissons, Gilbert and Snively, 2012. Locomotor forces and stress in the metapodia
of adult ostrich Struthio camelus and juvenile Albertosaurus sarcophagus
(Tyrannosauridae): Correlating anatomy, dynamics and finite element analysis.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 173.
Bell and Currie, 2014. Albertosaurus (Dinosauria: Theropoda) material
from an Edmontosaurus bonebed (Horseshoe Canyon Formation) near Edmonton:
Clarification of palaeogeographic distribution. Canadian Journal of Earth Sciences.
51(11), 1052-1057.
Torices, Reichel and Currie, 2014. Multivariate analysis of isolated tyrannosaurid
teeth from the Danek Bonebed, Horseshoe Canyon Formation, Alberta, Canada. Canadian
Journal of Earth Sciences. 51(11), 1045-1051.
Bell, Campione, Persons, Currie, Larson, Tanke and Bakker, 2017. Tyrannosauroid
integument reveals conflicting patterns of gigantism and feather evolution.
Biology Letters. 13: 20170092.
Funston, Powers, Whitebone, Brusatte, Scannella, Horner and Currie,
2020. Baby tyrannosaur bones from the Late Cretaceous of western North
America. The Society of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 147-148.
Albertosaurus? periculosus
Riabinin, 1930b
= Deinodon periculosus (Riabinin, 1930b) Kuhn, 1965
= Alectrosaurus periculosus (Riabinin, 1930b) Olshevsky 1991
= Jenghizkhan periculosus (Riabinin, 1930b) Olshevsky, 1995
= Tarbosaurus? periculosus (Riabinin, 1930b) Olshevsky, 1995
Late Maastrichtian, Late Cretaceous
Yuliangze Formation, Heilongjiang, China
Syntypes- (CCMGE unnumbered) tooth (47 x 23 x 9 mm), eight teeth, phalanx
Referred- teeth (Lu and Han, 2012)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Referred- (AEHM 1/789) lateral tooth (?x23.6x13.8 mm) (Bolotsky, 2011)
(AEHM 1/797) anterior maxillary tooth (?x12x11 mm) (Bolotsky, 2011)
(AEHM 1/799) anterior dentary tooth (?x14.3x10 mm) (Bolotsky, 2011)
(AEHM 1/800) lateral tooth (27.5x11.4x11.3 mm) (Bolotsky, 2011)
(AEHM 1/802) lateral tooth (42x16.3x9.7 mm) (Bolotsky, 2011)
(AEHM 1/804) (juvenile) posterior dentary tooth (24x12.2x7 mm) (Bolotsky, 2013)
(AEHM 1/805) (juvenile) posterior maxillary tooth (22.5x13x6.5 mm) (Bolotsky,
2013)
(AEHM 1/807) posterior maxillary tooth (22x12.5x6.5 mm) (Bolotsky, 2013)
(AEHM 1/808) (juvenile) anterior maxillary tooth (20.5x10.5x6 mm) (Bolotsky,
2013)
(AEHM 1/809) (juvenile) anterior maxillary tooth (18x10x6 mm) (Bolotsky, 2013)
(AEHM 1/810) posterior dentary tooth (13.5x?x? mm) (Bolotsky, 2013)
(AEHM 1/811) (juvenile) posterior dentary tooth (25x13.5x8 mm) (Bolotsky, 2013)
(AEHM 1/819) posterior dentary tooth (33x16.8x11.6 mm) (Bolotsky, 2011)
(AEHM 1/820) posterior dentary tooth (33x17.7x10.5 mm) (Bolotsky, 2011)
(AEHM 1/821) anterior maxillary tooth (48x18x10.2 mm) (Bolotsky, 2013)
(AEHM 1/822) tooth (40x19x11.8 mm) (Bolotsky, 2013)
(AEHM 1/823) posterior dentary tooth (38x17.9x11.5 mm) (Bolotsky, 2011)
(AEHM 1/824) posterior maxillary tooth (38.6x17.3x9.9 mm) (Bolotsky, 2011)
(AEHM 1/825) posterior maxillary tooth (?x17.2x9.9 mm) (Bolotsky, 2011)
(AEHM 1/826) posterior maxillary tooth (32x17x9.1 mm) (Bolotsky, 2011)
(AEHM 1/831) lateral tooth (?x19.5x12.2 mm) (Bolotsky, 2013)
(AEHM 1/834) posterior maxillary tooth (?x18.2x10 mm) (Bolotsky, 2013)
(AEHM 1/844) posterior dentary tooth (43x19.5x10 mm) (Bolotsky, 2013)
(AEHM 1/845) posterior dentary tooth (?x13.8x8 mm) (Bolotsky, 2013)
(AEHM 1/846) posterior dentary tooth (30x13.5x8.2 mm) (Bolotsky, 2011)
(AEHM 1/847) posterior dentary tooth (35.5x15.5x? mm) (Bolotsky, 2011)
(AEHM 1/848) posterior maxillary tooth (36x?x? mm) (Bolotsky, 2011)
(AEHM 1/849) anterior dentary tooth (29.5x13.6x9.5 mm) (Bolotsky, 2011)
(AEHM 1/850) posterior dentary tooth (29.3x16.3x10.9 mm) (Bolotsky, 2011)
(AEHM 1/851) posterior maxillary tooth (27.5x16.4x9 mm) (Bolotsky, 2011)
(AEHM 1/852) posterior dentary tooth (26.5x14.7x8 mm) (Bolotsky, 2011)
(AEHM 1/853) tooth (?x12.5x7.5 mm) (Bolotsky, 2013)
(AEHM 1/1004) incomplete ~fifteenth caudal vertebra (124 mm) (Bolotsky, 2013)
(AEHM 1/1005) incomplete ~twentieth caudal vertebra (122 mm) (Bolotsky, 2013)
(AEHM 1/1068) posterior dentary tooth (31x17x10 mm) (Bolotsky, 2013)
(AEHM 1/1070) anterior dentary tooth (26x12.5x7 mm) (Bolotsky, 2013)
(AEHM 1/1071) anterior dentary tooth (34.5x19.13.1 mm) (Bolotsky, 2013)
(AEHM 1/1073) posterior dentary tooth (?x16x8.8 mm) (Bolotsky, 2013)
(AEHM 1/1074) anterior maxillary tooth (22x11x10 mm) (Bolotsky, 2013)
(AEHM 1/1075) anterior maxillary tooth (26x13.5x8.2 mm) (Bolotsky, 2013)
(AEHM 1/1077) posterior maxillary tooth (32x15x8.5 mm) (Bolotsky, 2011)
(AEHM 1/1078) anterior maxillary tooth (27x12x8.4 mm) (Bolotsky, 2011)
(AEHM 1/1098) pedal phalanx II-2 (59 mm) (Bolotsky, 2013)
(AEHM 1/1100) pedal ungual II/IV (72.5 mm) (Bolotsky, 2013)
(AEHM 1/1106) mid caudal centrum (111 mm) (Bolotsky, 2013)
(AEHM 2/10) posterior dentary tooth (33.5x18.6x13.7 mm) (Bolotsky, 2011)
(AEHM 2/11) posterior maxillary tooth (37x19.3x12.2 mm) (Bolotsky, 2011)
(AEHM 2/12) anterior dentary tooth (40x18x11.5 mm) (Bolotsky, 2013)
(AEHM 2/13) posterior maxillary tooth (?x17.5x12.6 mm) (Bolotsky, 2011)
(AEHM 2/14) posterior dentary tooth (31.5x?x8.5 mm) (Bolotsky, 2013)
(AEHM 2/424) posterior maxillary tooth (?x19.2x12 mm) (Bolotsky, 2011)
(AEHM 2/425) posterior dentary tooth (?x20x12.2 mm) (Bolotsky, 2013)
(AEHM 2/426) posterior dentary tooth (?x?x11 mm) (Bolotsky, 2013)
(AEHM 2/427) (juvenile) posterior dentary tooth (16.5x9.8x6.7 mm) (Bolotsky,
2011)
(AEHM 2/428) premaxillary tooth (26 mm) (Bolotsky, 2011)
(AEHM 2/431) (juvenile) anterior maxillary tooth (16.5x9.2x6.2 mm) (Bolotsky,
2011)
(AEHM 2/434) posterior dentary tooth (?x20.7x12.5 mm) (Bolotsky, 2011)
(AEHM 2/435) posterior dentary tooth (36.5x12.9x11.5 mm) (Bolotsky, 2011)
(AEHM 2/436) anterior dentary tooth (?x15.4x10.3 mm) (Bolotsky, 2011)
(AEHM 2/925) proximal tibia (Bolotsky, 2013)
(AEHM 2/1027) posterior maxillary tooth (?x11.4x9.8 mm) (Bolotsky, 2011)
(AEHM 2/1028) posterior maxillary tooth (24x14.5x7.3 mm) (Bolotsky, 2011)
(AEHM 2/1038) posterior maxillary tooth tooth (46x17.9x11.2 mm) (Bolotsky, 2011)
?(AEHM 2/1042) tooth (Bolotsky, 2013)
(AEHM 2/1044) pedal ungual II (69 mm) (Bolotsky, 2013)
(AEHM 2/1114) (juvenile) posterior maxillary tooth (20x11x5.5 mm) (Bolotsky,
2013)
?(AEHM 2/coll.) fourth cervical vertebra (Alifanov and Bolotsky, 2002)
Comments- The holotype's repository (verified by Averianov, pers. comm.
2015) is based on the recent redescription of 'Aspideretes' planicostatus
(as Amuremys planicostata) by Danilov et al. (2002), which was collected
by Riabinin in the same locality as periculosus and described by him
in the same paper. Riabinin based the species on nine teeth with a possibly
referred phalanx, and provided the dimensions of the largest. Thus the periculosus
material consists of syntypes, of which the sole figured specimen should be
designated the lectotype once redescribed. These teeth were discovered in 1916-1917
and first mentioned by Riabinin (1930a) as Dryptosaurus? sp. before being
described by him (1930b) as a species of Albertosaurus, based on smaller
size than Tyrannosaurus and supposedly Gorgosaurus. Every generic
reassignment since has been done in taxonomic lists without justification, except
possibly Olshevsky's (1995) which has not been translated from Japanese. Lu
and Han (2012) referred several teeth from the same locality to not only tyrannosaurids,
but carcharodontosaurids, Fukuiraptor, carcharodontosaurids and a possible
new theropod. Bolotsky (2013) thought these could all be tyrannosaurid however.
Notably, the Yuliangze Formation continues across the Amur River into Russia
as the Udurchukan Formation of the Tsagayan Group, which has preserved numerous
tyrannosaurid remains. Alifanov and Bolotsky (2002) refer some of these Russian
teeth to periculosus and Aublysodon, which were later briefly
described as Tyrannosauridae indet. by Bolotsky (2011). Bolotsky (2013) described
Udurchukan tyrannosaurid material in depth in his thesis, comparing them to
Yuliangze teeth as well. He found the smaller Udurchukan and Yuliangze teeth
are comparable and that they are more similar to albertosaurines than tyrannosaurines.
While Bolotsky didn't assign particular teeth to each subfamily, those which
are within his small range are assigned to periculosus here, though there
is some size overlap. All postcrania (except a metacarpal I) were said to be
albertosaurine-sized, and the cervical mentioned by Alifanov and Bolotsky (2002)
was said to be from a ~6-8 meter long individual. It's possible at least some
of these are from juvenile tyrannosaurines, Alioramus, or more basal
tyrannosauroids.
References- Riabinin, 1930a. Mandschurosaurus amurensis nov. gen.
nov. sp., a hadrosaurian dinosaur from the Upper Cretaceous of Amur River. Memoir
of Russian Paleontological Society. 2, 36 pp.
Riabinin, 1930b. On the age and fauna of the dinosaur beds on the Amur River.
Zapiski Rossiiskogo minyeralogichyeskogo obshchyestva [Memoirs of the Russian
Mineralogical Society], second series. 59(1), 41-51.
Kuhn, 1965. Saurischia {Supplementum I}. Fossilium Catalogus I Animalia Pars
109. 94 pp.
Bolotsky and Moiseenko, 1988. O Dinozavrakh Priamur'ya. Doklady Akademii Nauk.
SSSR, Dal'nevostkhnoe Otdelenie Amursky Kompleksny Naukno-Issledovatelsky Institut,
Blagoveshensk. 37 pp.
Olshevsky, 1991. A Revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku
Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.
Alifanov and Bolotsky, 2002. New data about the assemblages of the Upper Cretaceous
carnivorous dinosaurs (Theropoda) from the Amur region. In Kirillova (ed.).
Fourth International Symposium of IGCP 434. Cretaceous continental margin of
East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
Danilov, Bolotsky, Averianov and Donchenko, 2002. A new genus of lindholmemydid
turtle (Testudines: Testudinoidea) from the Late Cretaceous of the Amur region,
Russia. Russian Journal of Herpetology. 9(2), 155-168.
Bolotsky and Bolotsky, 2008. Tyrannosaurids from the Amur (Heilongjiang) river
basin. Abstracts of the International Dinosaur Symposium in Fukui 2008. 19-20.
Bolotsky, 2009. Tyrannosaurid teeth from Maastricht of Amur Region / / Modern
paleontology: Classical and new methods. Moscow: PIN RAS. 83-88.
Bolotsky, 2011. On paleoecology of carnivorous dinosaurs (Tyrannosauridae, Dromaeosauridae)
from Late Cretaceous fossil deposits of Amur region, Russian far East. Global
Geology. 14(1), 1-6.
Lu and Han, 2012. The discovery of Late Cretaceous theropod dinosaur teeth from
Jiayin Area, Heilongjiang Province and its significance. Acta Geologica Sinica.
86(3), 363-270.
Bolotsky, 2013. Tyrannosaurid dinosaurs (Coelurosauria) from Upper Cretaceous
of Amur/Heilongjiang Area. Masters thesis, Jilin University. 73 pp.
Tyrannosaurinae Osborn, 1906 sensu Matthew
and Brown, 1922
Definition- (Tyrannosaurus rex <- Albertosaurus sarcophagus)
(Holtz, 2004; modified from Currie et al., 2003)
Other definitions- (Tyrannosaurus rex <- Albertosaurus sarcophagus,
Daspletosaurus torosus, Gorgosaurus libratus) (modified from Sereno, 1998)
(Tyrannosaurus rex <- Aublysodon mirandus) (modified from Holtz,
2001)
(Tyrannosaurus rex <- Gorgosaurus libratus, Albertosaurus
sarcophagus) (Sereno, in prep.)
= Shanshanosaurinae Dong, 1977 sensu Olshevsky, 1995
= Tyrannosaurinae sensu Sereno in prep.
Definition- (Tyrannosaurus rex <- Gorgosaurus libratus, Albertosaurus
sarcophagus)
Diagnosis- (after Carr, 2005) dorsal surface of dorsotemporal fossa of
squamosal is convex; nasal process of frontal elongate; nasal process of frontal
narrow; sigittal crest of frontal tall and long.
Comments- Sereno's (in prep.) definition is a revision of Currie et al.'s
(2003), adding Gorgosaurus as an external specifier. It does do a better
job at maintaining stability if albertosaurines are paraphyletic. And since
Tyrannosauridae has multiple internal specifiers, this isn't part of a node-stem
triplet, so I tentatively agree with Sereno.
References- Osborn, 1906. Tyrannosaurus, Upper Cretaceous carnivorous
dinosaur (Second communication). Bulletin of the American Museum of Natural
History. 22(16), 281-296.
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.
Dong, 1977. On the dinosaurian remains from Turpan, Xinjiang. Vertebrata PalAsiatica.
15(1), 59-66.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku
Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210(1), 41-83.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. In Tanke and
Carpenter (eds.). Mesozoic Vertebrate Life. 64-83.
Currie, Hurum and Sabath, 2003. Skull structure and evolution in tyrannosaurid
dinosaurs. Acta Palaeontologica Polonica. 48(2), 227-234.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Carr, 2013. Using ontogeny and phylogeny to test hypotheses of anagenesis in
the vertebrate fossil record: A case study of the sister group relationship
between Daspletosaurus and Tyrannosaurus (Dinosauria, Coelurosauria).
Journal of Vertebrate Paleontology. Program and Abstracts 2013, 101.
unnamed tyrannosaurine 'Aguja tyrannosaur' (Lehman, 1985)
Late Campanian, Late Cretaceous
Upper Aguja Formation, Texas, US
Material- (LSUMG 489:5580) tooth fragment (Sankey, 2001)
(LSUMG 489 coll.) tooth (Sankey, 2008)
(LSUMG 491 coll.) four teeth (Sankey, 2008)
(LSUMG V-1312; Morphotype C of Standhardt) tooth fragment (Standhardt, 1986)
(LSUMG V-1313; Morphotype B of Standhardt) partial tooth (Standhardt, 1986)
(TMM 40573-1) tibia (610 mm) (Lehman and Wick, 2013)
(TMM 41918-24) (juvenile) fragmentary frontal (Lehman and Wick, 2013)
(TMM 42533-5) pedal phalanx IV-3/4 (Lehman and Wick, 2013)
(TMM 42534-15) tooth (Lehman, 1989)
(TMM 42545-7) pedal phalanx IV-3, phalanx IV-4 (Lehman and Wick, 2013)
(TMM 42880-7) dentary tooth (Lehman and Wick, 2013)
(TMM 42880-8) (juvenile) premaxillary tooth (Lehman and Wick, 2013)
(TMM 43057-142) pedal phalanx III-1 (Lehman and Wick, 2013)
(TMM 43057-505) ~twenty-first to twenty-fourth caudal vertebra (Lehman and Wick, 2013)
(TMM 45905-1) frontal (Lehman and Wick, 2013)
(TMM 45906-1) (19 year old, ~650 kg adult) (femur ~765 mm) incomplete
metatarsal II, phalanx II-1, metatarsal III (488 mm), metatarsal IV
(456 mm) (Lehman and Wick, 2013)
(TMM 45907-1) pedal ungual II (~120 mm) (Lehman and Wick, 2013)
(TMM 45908-1) pedal phalanx IV-3/4 (Lehman and Wick, 2013)
(TMM 45908-2) dentary tooth (Lehman and Wick, 2013)
(TMM 45908-3) anterior maxillary tooth (Lehman and Wick, 2013)
(TMM 45909-1) ~thirtieth-thirty-third caudal vertebra (Lehman and Wick, 2013)
(TMM 45910-1) incomplete astragalus (Lehman and Wick, 2013)
(TMM coll.) many partial teeth (Lehman and Wick, 2013)
Late Campanian, Late Cretaceous
Aguja Formation Mexico
Material- teeth, limb elements (Westgate, Pittman, Brown and Cope, 2002)
material (Montellano, Monroy, Hernandez-Rivera and Torres, 2009)
? tooth (Rivera-Sylva, Hedrick, Guzman-Gutierrez, Gonzalez and Dodson, 2011)
Comments-
Lehman (1985) originally identified tyrannosaurid teeth from the Aguja
Formation of Texas in his thesis. Standhardt (1986) identified
three partial teeth from there as carnosaurs in his thesis, of which
his Morphotype A (LSUMG V-1375) is here excluded as it lacks mesial
serrations so is more likely dromaeosaurid. A large incomplete
femur (LSUMG V-1226) referred to Carnosauria incertae sedis is more
likely hadrosaurid based on the lack of an ectocondylar tuber.
Langston et al. (1989) identified Daspletosaurus
remains from the lower and upper beds of the Aguja Formation of
Texas. Rowe et al. (1992) stated "at least one large
tyrannosaurid" (TMM coll.) was present at the Terlingua site.
Rowe et al. (1992) identified cf. Dromaeosaurus teeth
(including TMM 43057-314) from the Aguja Formation of Texas. Sankey (1998) later
identified Dromaeosaurus teeth from another area of that formation, but
these and Rowe et al.'s specimens were referred to Theropoda "family and
genus undetermined." They consisted of two tooth fragments (LSUMG 5483
and 6239) which were similar to Dromaeosaurus except in lacking a lingually
twisted mesial carina, and were reidentified as tyrannosaurid teeth by Sankey
et al. (2005).
Lehman and Wick (2013) describe tyrannosaurid remains from the upper
Aguja Formation as the 'Aguja tyrannosaur', stating "although we
consider it unlikely, it is possible that the specimens could pertain
to more than one taxon." They note premaxillary tooth
TMM 42880-8 lacks serrations, lateral teeth TMM 42880-7 and 45908-2
have labiolingual compression similar to albertosaurines, a high
frontal parietal crest like tyrannosaurines, and gracile hindlimb
elements as in Appalachiosaurus
and juvenile albertosaurines. The frontal morphology is
diagnostic, but the authors err on the side of caution and leave it
unnamed given the controversial taxonomic history of fragmentary
tyrannosauroid specimens.
From Mexico, Westgate et al. (2002) reported "tyrannosaurid ... teeth
and limb elements" from Chihuahua, Montellano et al. (2009) cited "a
form of an indetermined tyrannosaurid" from Coahuila and Rivera-Sylva
et al. (2011) reported "one possible tyrannosaurid tooth" from a new
locality in Coahuila.
References- Lehman, 1985. Stratigraphy, sedimentology, and paleontology
of Upper Cretaceous (Campanian-Maastrichtian) sedimentary rocks in Trans-Pecos,
Texas. PhD thesis, University of Texas at Austin. 299 pp.
Standhardt, 1986. Vertebrate paleontology of the Cretaceous/Tertiary transition
of Big Bend National Park, Texas. PhD thesis, Louisiana State University. 298
pp.
Lehman, 1989. Chasmosaurus mariscalensis, sp. nov., a new ceratopsian
dinosaur from Texas. Journal of Vertebrate Paleontology. 9(2), 137-162.
Langston, Standhardt and Stevens, 1989. Fossil vertebrate
collecting in the Big Bend - history and perspective. In Busbey and Lehman (eds.).
Vertebrate paleontology, biostratigraphy, and depositional environments, Latest
Cretaceous and Tertiary, Big Bend area, Texas. SVP 1989 Guidebook. 11-21.
Rowe, Ciffelli, Lehman and Weil, 1992. The Campanian Terlingua local fauna,
with a summary of other vertebrates from the Aguja Formation, Trans-Pecos, Texas.
Journal of Vertebrate Paleontology. 12(4), 472-493.
Sankey, 1998. Vertebrate paleontology and magnetostratigraphy of the upper Aguja
Formation (Late Campanian), Talley Mountain area, Big Bend National Park, Texas.
PhD thesis, Louisiana State University. 263 pp.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend,
Texas. Journal of Paleontology. 75(1), 208-215.
Westgate, Pittman, Brown and Cope, 2002. Continued excavation of the first dinosaur
community from Chihuahua, Mexico. Journal of Vertebrate Paleontology. 22(3),
118A.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous
(Campanian-Maastrichtian), Big Bend National Park, Texas. In Carpenter (ed.).
The Carnivorous Dinosaurs. 127-152.
Montellano, Monroy, Hernandez-Rivera and Torres, 2009. Late Cretaceous microvertebrate
fauna from the northern state of Coahuila, Mexico. Journal of Vertebrate Paleontology.
29(3), 151A.
Rivera-Sylva, Hedrick, Guzman-Gutierrez, Gonzalez and Dodson, 2011. A new Campanian
vertebrate locality from northwestern Coahuila, Mexico. Journal of Vertebrate
Paleontology. Program and Abstracts 2011, 179.
Lehman and Wick, 2013. Tyrannosauroid dinosaurs from the Aguja
Formation (Upper Cretaceous) of Big Bend National Park, Texas. Earth
and Environmental Science Transactions of the Royal Society of
Edinburgh. 103, 1-15.
unnamed Tyrannosaurinae (Alifanov and Bolotsky, 2002)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- (AEHM 1/790) lateral tooth (71x?x14.4 mm) (Bolotsky, 2011)
(AEHM 1/791) lateral tooth (64.9x?x13 mm) (Bolotsky, 2011)
(AEHM 1/798) premaxillary tooth (?x14x13 mm) (Bolotsky, 2013)
(AEHM 1/803) posterior dentary tooth (61x21.5x18.5 mm) (Bolotsky, 2013)
(AEHM 1/812) posterior maxillary tooth (60x?x? mm) (Bolotsky, 2013)
(AEHM 1/813) anterior maxillary tooth (57x?x15.2 mm) (Bolotsky, 2013)
(AEHM 1/818) anterior maxillary tooth (56x20x? mm) (Bolotsky, 2013)
(AEHM 1/832) lateral tooth (54x?x13 mm) (Bolotsky, 2013)
(AEHM 1/841) posterior maxillary tooth (73x26.7x17.5 mm) (Bolotsky, 2011)
(AEHM 1/842) posterior dentary tooth (59x26.4x21 mm) (Bolotsky, 2011)
(AEHM 1/843) posterior maxillary tooth (53x24.8x14 mm) (Bolotsky, 2013)
(AEHM 1/1072) posterior maxillary tooth (58x?x? mm) (Bolotsky, 2013)
(AEHM 1/1099) metacarpal I (68 mm) (Bolotsky, 2013)
(AEHM 1/1657) posterior maxillary tooth (81x31x20 mm) (Bolotsky, 2013)
(AEHM 2/421) posterior maxillary tooth (72x27.9x16.5 mm) (Bolotsky, 2011)
(AEHM 2/422) posterior maxillary tooth (58x22.2x12 mm) (Bolotsky, 2013)
(AEHM 2/423) posterior dentary tooth (?x21.5x13.5 mm) (Bolotsky, 2013)
(AEHM 2/1037) anterior maxillary tooth (73x26.5x18 mm) (Bolotsky, 2011)
(AEHM 2/1041) posterior maxillary tooth (?x27x17 mm) (Bolotsky, 2013)
Comments- These were referred to cf. Tarbosaurus sp. by Alifanov
and Bolotsky (2002), and briefly described as Tyrannosauridae indet. by Bolotsky
(2011). Bolotsky (2013) analyzed the remains in detail and proposed the large
teeth and metacarpal I belonged to a tyrannosaurine, as opposed to the smaller
material which is albertosaurine and here provisionally referred to Albertosaurus?
periculosus. While Bolotsky never assigned specific teeth to Tyrannosaurinae,
I've placed those larger than his listed ranges for albertosaurine teeth here,
but there is some overlap.
References- Alifanov and Bolotsky, 2002. New data about the assemblages
of the Upper Cretaceous carnivorous dinosaurs (Theropoda) from the Amur region.
In Kirillova (ed.). Fourth International Symposium of IGCP 434. Cretaceous continental
margin of East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
Bolotsky, 2011. On paleoecology of carnivorous dinosaurs (Tyrannosauridae, Dromaeosauridae)
from Late Cretaceous fossil deposits of Amur region, Russian far East. Global
Geology. 14(1), 1-6.
Bolotsky, 2013. Tyrannosaurid dinosaurs (Coelurosauria) from Upper Cretaceous
of Amur/Heilongjiang Area. Masters thesis, Jilin University. 73 pp.
Alioramini Olshevsky, 1995
Definition- (Alioramus remotus <- Tyrannosaurus rex, Albertosaurus
sarcophagus, Proceratosaurus bradleyi) (after Lu et al., 2014)
References- Olshevsky, 1995. The origin and evolution of the tyrannosaurids.
Kyoryugaku Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.
Lu, Yi, Brusatte, Yang, Li and Chen, 2014. A new clade of Asian Late Cretaceous
long-snouted tyrannosaurids. Nature Communications. 5, 3788.
Alioramus Kurzanov, 1976
= Qianzhousaurus Lu, Yi, Brusatte, Yang, Li and Chen, 2014
A. remotus Kurzanov, 1976
= Alioramus altai Brusatte, Carr, Erickson, Bever and Norell, 2009
= Qianzhousaurus sinensis Lu, Yi, Brusatte, Yang, Li and Chen, 2014
= Alioramus sinensis (Lu, Yi, Brusatte, Yang, Li and Chen, 2014) Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017
Maastrichtian, Late Cretaceous
Nogon Tsav, Nemegt Formation, Mongolia
Holotype- (GIN 3141/1) (juvenile) incomplete skull (~700 mm), mandible,
four cervical vertebrae, partial tibia, proximal fibula, pedal ungual I, distal
metatarsal II, phalanx II-1, pedal ungual II, distal metatarsal III, phalanx
III-1, pedal ungual III, distal metatarsal IV, phalanx IV-1, pedal ungual IV
Early Maastrichtian, Late Cretaceous
Tsagan Khushuu, Nemegt Formation, Mongolia
Referred- (IGM 100/1844; holotype of Alioramus altai) (9 year
old juvenile; 369 kg) incomplete skull (~635 mm), mandibles (one partial), hyoids,
atlantal intercentrum, altantal neurapophyses, incomplete axis (36 mm), incomplete
third cervical vertebra (42 mm), incomplete fourth cervical vertebra (42 mm),
incomplete fifth cervical vertebra (65 mm), incomplete sixth cervical vertebra
(75 mm), seventh cervical vertebra (57 mm), eighth cervical vertebra (60 mm),
ninth cervical vertebra (67 mm), tenth cervical vertebra (51 mm), seven cervical
ribs, partial anterior dorsal vertebra (55 mm), posterior dorsal vertebra (55
mm), dorsal vertebral fragment, two dorsal ribs, incomplete sacrum (?,?,75,79,97
mm), fourth sacral rib, fifth sacral rib, proximal caudal vertebra (87 mm),
proximal caudal vertebra (82 mm), distal caudal vertebra (84 mm), mid chevron,
incomplete ilium, ischia (430 mm; one partial), femora (560 mm; one fragmentary),
distal tibia (101 mm transverse width), distal fibula, astragalus, calcaneum,
distal tarsal III, distal tarsal IV, partial metatarsal I, phalanges I-1 (48
mm), pedal ungual I (36 mm), proximal metatarsal II, partial metatarsals III,
phalanx III-1, proximal metatarsal IV, metatarsals V (one incomplete, one partial),
metatarsal fragments, several pedal phalanges (Brusatte et al., 2009)
Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
(GM F10004; holotype of Qianzhousaurus sinensis) incomplete skull (900
mm), incomplete mandible, atlantal intercentrum (19.4 mm), axis (54.5 mm), third
cervical vertebra (48.8 mm), fourth cervical vertebra (55.8 mm), sixth cervical
vertebra (86.7 mm), seventh cervical vertebra (88.1 mm), eighth cervical vertebra
(91.5 mm), ninth cervical vertebra (85.1 mm), tenth cervical vertebra (78.9
mm), first dorsal vertebra (65.5 mm), second dorsal vertebra (68.4 mm), third
dorsal vertebra (69.8 mm), fourth dorsal vertebra (75.1 mm), partial ~third
caudal centrum, ~fourth caudal vertebra (85.1 mm), ~fifth caudal vertebra (96.1
mm), ~sixth caudal vertebra (94.1 mm), ~seventh caudal vertebra (94.1 mm), ~eighth
caudal vertebra (98.5 mm), partial ~ninth caudal vertebra, partial ~nineteeth
caudal vertebra, ~twentieth caudal vertebra (96.7 mm), ~twenty-first caudal
vertebra (102.7 mm), ~twenty-second caudal vertebra (100.9 mm), ~twenty-third
caudal vertebra (97.5 mm), ~twenty-fourth caudal vertebra (97.6 mm), ~twenty-fifth
caudal vertebra (88.3 mm), ~twenty-sixth caudal vertebra (84.7 mm), ~twenty-seventh
caudal vertebra (80.4 mm), ~twenty-eighth caudal vertebra (75.2 mm), ~twenty-ninth
caudal vertebra (68.3 mm), scapulocoracoids (one partial), partial ilia, incomplete
femur (700 mm), tibia (760 mm), partial fibula, astragalus, calcaneum, metatarsal
I (75 mm), incomplete metatarsal III, incomplete metatarsal IV (Lu, Yi, Brusatte,
Yang, Li and Chen, 2014)
Diagnosis- (after Kurzanov, 1976) elongate skull (length/height ratio
>3); 16-17 maxillary teeth (ontogenetic?); 18-20 dentary teeth (ontogenetic?).
(after Brusatte et al., 2009 for A. altai) accessory pneumatic fenestra
posterodorsal to promaxillary fenestra of maxilla (ontogenetic?); maxillary
fenestra enlarged and 1.9 times longer than deep; laterally projecting jugal
horn; thick ridge on dorsal surface of the ectopterygoid; anteroposteriorly
elongate anterior mylohyoid foramen of splenial; thin epipophysis on atlantal
neurapophysis that terminates at a sharp point; pneumatic pocket on anterior
surface of cervical transverse processes (ontogenetic?); external pneumatic
foramina on dorsal ribs (ontogenetic?); anterodorsally inclined midline ridge
on the lateral surface of the ilium.
Other diagnoses- Kurzanov (1976) listed many additional characters, most
of which are probably due to the Alioramus type's juvenile age- 'average'
size; elongate snout; series of prominent nasal rugosities; small postorbital
boss; labiolingually compressed teeth. Two rows of maxillary nutrient foramina
are present in most tyrannosaurids (Currie, 2003), as are the laterosphenoid
contacts noted by Kurzanov (forms part of the supratemporal cavity and contacts
the postorbital). Currie also noted the trigeminal foramen near certainly contacted
the laterosphenoid as opposed to being completely contained by the prootic.
While he defended the prominence of the nasal rugostities as potentially diagnostic,
they are lower in IGM 100/1844.
Brusatte et al. (2009) stated several characters united the Alioramus
specimens in their analysis, most being previously used by Kurzanov except for
the long posterior squamosal process. Yet Carr (2005) notes that juvenile Tyrannosaurus
have long processes, making this potentially ontogenetic. Among characters listed
in the diagnosis for A. altai which are unknown in the A. remotus
holotype, the palatine pneumatic recess extends posteriorly beyond the posterior
margin of the vomeropterygoid process in juvenile Daspletosaurus and
Tyrannosaurus more than in adults.
Comments- The holotype of Alioramus remotus
was found between 1969-1973 and described by Kurzanov (1976) as a form
of primitive tyrannosaurid. Currie et al. (2003) found Alioramus to be the sister
taxon of Tarbosaurus because they both lack a lacrimal process on the
nasal, though this is present in Daspletosaurus as well. In addition,
Hurum and Sabath (2003) note Alioramus and Tarbosaurus share a
dentary-angular interlocking mechanism which makes the mandible rigid. Currie
(2003) suggested the specimen could be a juvenile Tarbosaurus based on
skull proportions and juvenile characters. He stated the prominent nasal rugosities
and high tooth count argue against this, but juvenile Tyrannosaurus have
high tooth counts and some juvenile Daspletosaurus and Tarbosaurus
have rows of nasal rugosities, albeit lower ones as in the A. altai and
Qianzhousaurus holotypes. Holtz (2004) recovered Alioramus in
two possible positions- just basal to Tyrannosauridae or sister to Tarbosaurus
+ Tyrannosaurus. The former position is due to the high tooth count,
low snout and slender dentary, which are all possible juvenile characters. The
latter position was due to the thick parietal nuchal crest, reduced basal tubera,
and posteroventrally directed occipital region. Carr (2005) recovered Alioramus
in an uncertain position basal to Tyrannosauridae, but this could be due to
juvenile characters. However, the evidence cannot be examined as characters
excluding the taxon from Tyrannosauridae were not given, nor was Alioramus
included in the printed data matrix. Brusatte et al. (2009) found Alioramus
to be a basal tyrannosaurine using an updated version of Carr's matrix, but
importantly coded it as if it were an adult when both morphology and histology
show known specimens are juveniles. Thus its position is suspect, as similarly
aged Tyrannosaurus individuals also emerge as basal tyrannosaurines if
run in a similar matrix (Carr, 2005). The same could be said of Lu et al.'s
(2014) analyses adding Qianzhousaurus, where alioramins emerge either
as basal tyrannosaurines or just basal to Tyrannosauridae, and Brusatte and
Carr (2016) who found alioramins to be the most basal tyrannosaurines. IGM 100/1844
also provides further evidence for a relationship to Tarbosaurus, as
it has a subcutaneous flange on the maxilla and a deep pneumatic fossa on the
dorsal surface of the posterior centrodiapophyseal lamina, both otherwise only
known in that genus. However, they also noted additional characters which differ
between Alioramus and Tarbosaurus of the same size (ZPAL MgD-I/29,
31 and 175)- shallow maxilla and dentary; maxilla less convex ventrally; smaller
postorbital boss (not in the Qianzhousaurus type); postorbital lacks
an orbital process; more dentary teeth; muscular fossa above surangular foramen
faces mostly dorsally; laterally projecting jugal horn (not in the Qianzhousaurus
type); deep pocket behind surangular fenestra; fibular facet of tibia faces
strongly laterally; lateral malleolus of tibia projects less distolaterally.
The first six characters are typical of juveniles and could potentially indicate
Alioramus individuals are larger at a younger age than ZPAL MgD-I/29
and 31, or that different individuals acquire adult features at different ages.
The jugal horn and surangular pocket are ornamental and pneumatic features respectively,
which show a high amount of individual variation. Brusatte et al. even state
that an ontogenetic decrease in pneumaticity is known in theropods and that
Tarbosaurus itself is known to lose pneumatic vertebral features with
age, potentially explaining the surangular pocket and some of A. altai's
supposed diagnostic features (see diagnosis above). Ontogenetic variation in
tyrannosaurid tibiae has not been examined yet. While Brusatte et al. claimed
that ornamentation increases in ontogeny in dinosaurs, this is not always the
case as shown by juvenile tyrannosaurines with larger nasal rugosities and the
newly discovered ontogenetic changes in Triceratops (?= Torosaurus)
and Pachycephalosaurus (= Stygimoloch and Dracorex). Despite
the fact most differences could be explained by ontogeny and the unique similarities
present in Alioramus and the contemporaneous Tarbosaurus, the
recently discovered specimen named as Qianzhousaurus is as large as several
traditional Tarbosaurus specimens yet retains an Alioramus morphology.
This would seem to indicate the taxa are distinct after all.
Alioramus altai- Originally announced in abstracts by Bever et
al. (2009) and Norell et al. (2009), Brusatte et al. (2009) erected the species
Alioramus altai based on a partial skeleton discovered in 2001 from Tsagan Khushuu. However, the listed diagnostic characters
are problematic. Most are not determinable in A. remotus (accessory pneumatic
fenestra posterodorsal to promaxillary fenestra of maxilla; maxillary fenestra
enlarged and 1.9 times longer than deep; thick ridge on dorsal surface of the
ectopterygoid; palatine pneumatic recess extending posteriorly beyond posterior
margin of vomeropterygoid process [also in Daspletosaurus sp.]; thin
epipophysis on atlantal neurapophysis that terminates at a sharp point; external
pneumatic foramina on dorsal ribs; anterodorsally inclined midline ridge on
the lateral surface of the ilium [also in some Gorgosaurus, Daspletosaurus
and Tyrannosaurus specimens]) or potentially determinable but unreported
(anteroposteriorly elongate anterior mylohyoid foramen of splenial; pneumatic
pocket on anterior surface of cervical transverse processes). The laterally
projecting jugal horn was also coded as present in A. remotus, and Brusatte
et al. (2012) note it may be present in that species based on Kurzanov's description.
Having 20 dentary teeth instead of 18 is within the range of variation in other
tyrannosaurid species. The subcutaneous flange on the maxilla is known to vary
in Tarbosaurus. The authors themselves note in the supplementary information
that some of the characters they list as distinguishing A. altai from
A. remotus vary within other tyrannosaurid species- anterior process
of quadratojugal terminates posterior to anterior margin of lateral temporal
fenestra; squamosal anterior process extends anterior to anterior margin of
lateral temporal fenestra; epipterygoid not bifurcated ventrally (which may
be due to damage in A. remotus). The number and prominence of nasal rugosities
is highly variable in tyrannosaurids, so A. remotus having six large
rugosities is not significant compared to A. altai's three low ones.
Finally, Brusatte et al. list three characters which are size-related in other
tyrannosaurid taxa- 17 maxillary teeth instead of 16; single dorsoventral groove
between basal tubera; tapering anterior process of the parietals overlapping
frontals on the midline. They considered these potentially diagnostic since
the holotypes are similar in size, but at least the maxillary tooth count and
parietal anterior process morphology are variable in similar-sized specimens.
Here the basal tubera groove is considered individual variation as well, as
this has only been noted to be ontogenetic in Tyrannosaurus. Brusatte
et al. (2012) added two more characters, which are both unknown in A. remotus
as well- dorsally extending and conical lacrimal horn; ventrally sloping posterior
ilial margin. Those authors also noted the anteroventrally sloping dorsal quadratojugal
margin differs from the horizontal margin of A. remotus, but found this
was also ontogenetic in Tyrannosaurus. Oddly, though Brusatte et al.
(2012) conclude almost all of their proposed A. altai autapomorphies
cannot be evaluated for A. remotus, are ontogenetically and/or individually
variable in other tyrannosaurids, they still retain it as a separate species.
However, Brusatte (2013) notes "it is possible, and perhaps probable, that
future work on the A. remotus holotype" in progress by Alifanov,
will show the species are synonymous. As of Carr et al. (2017), the authors state "we take the view that Alioramus altai is synonymous with A. remotus."
Qianzhousaurus sinensis- Lu et al. (2014) describe a new partial
skeleton as Qianzhousaurus sinensis, finding it to be in a trichotomy
with Alioramus remotus and A. altai. They call this clade Alioramini.
Most described differences are acknowledged to be ontogenetic, expected as Qianzhousaurus
is ~25% larger than A. altai and A. remotus. Among the supposed
diagnostic characters, the "extremely reduced premaxilla (maximum anteroposterior
length of the main body of the bone is ~2.2% of the total basal skull length,
...)" cannot be coded in Alioramus remotus or A. altai as
those specimens don't preserve premaxillae. The fenestrated maxillary pneumatic
excavation in the ascending process cannot be coded in A. remotus, but
as A. altai expresses this as a fossa and details of pneumatic features
are highly variable between individuals, the differences in size, shape and
placement between these specimens ("larger, located further posteriorly,
oriented nearly vertically instead of horizontally" in Qianzhousaurus)
is not considered taxonomically important. Finally, Qianzhousaurus lacks
a vertical ilial ridge unlike A. altai, which is again unknown in A.
remotus. This in itself is here considered insufficient to diagnose a new
taxon, especially as it fails to distinguish Qianzhousaurus from Alioramus
remotus, and all three specimens are from stratigraphically equivalent horizons.
Lu et al. further use geographic distance to distinguish their genus but this
is surely inconsequential. Given the above, Qianzhousaurus sinensis is
a junior synonym of Alioramus remotus.
References- Kurzanov, 1976. A new Late Cretaceous carnosaur from Nogon-Tsav
Mongolia. Sovmestnaa Sovetsko-Mongolskaa Paleontologiceskaa Ekspeditcia, Trudy.
3, 93-104.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous
of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Currie, Hurum and Sabath, 2003. Skull structure and evolution in tyrannosaurid
dinosaurs. Acta Palaeontologica Polonica. 48(2), 227-234.
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.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Bever, Brusatte, Carr and Norell, 2009. The braincase of a new tyrannosaurid
from the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 29(3),
63A.
Brusatte, Carr, Erickson, Bever and Norell, 2009. A long-snouted, multihorned
tyrannosaurid from the Late Cretaceous of Mongolia. Proceedings of the National
Academy of Sciences. 106(41), 17261-17266.
Norell, Brusatte, Carr, Bever and Erickson, 2009. A remarkable long-snouted,
multi-horned tyrannosaurid from the Late Cretaceous of Mongolia. Journal of
Vertebrate Paleontology. 29(3), 155A.
Bever, Brusatte, Balanoff and Norell, 2011 online. Alioramus altai, Digital Morphology. http://digimorph.org/specimens/Alioramus_altai/
Bever, Brusatte, Balanoff and Norell, 2011. Variation, variability, and the
origin of the avian endocranium: Insights from the anatomy of Alioramus altai
(Theropoda: Tyrannosauroidea). PLoS ONE. 6(8), e23393.
Brusatte, Carr and Norell, 2012. The osteology of Alioramus, a gracile
and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous
of Mongolia. Bulletin of the American Museum of Natural History. 366, 197 pp.
Gold, Brusatte and Norell, 2012. Pneumaticity patterns in the skull of Alioramus
altai, a long-snouted tyrannosaurid (Dinosauria: Theropoda), from the Late
Cretaceous of Mongolia. Journal of Vertebrate Paleontology. Program and Abstracts
2012, 102.
Brusatte, 2013. The phylogeny of basal coelurosaurian theropods (Archosauria:
Dinosauria) and patterns of morphological evolution during the dinosaur-bird
transition. PhD thesis. Columbia University. 944 pp.
Brusatte, Lu, Carr, Williamson and Norell, 2014. A clade of long-snouted tyrannosaurids
ranged across Asia during the Latest Cretaceous. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 99.
Lu, Yi, Brusatte, Yang, Li and Chen, 2014. A new clade of Asian Late Cretaceous
long-snouted tyrannosaurids. Nature Communications. 5, 3788.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017. A new tyrannosaur
with evidence for anagenesis and crocodile-like facial sensory system.
Scientific Reports. 7:44942.
Dynamoterror McDonald, Wolfe and Dooley Jr, 2018
D. dynastes McDonald, Wolfe and Dooley Jr, 2018
Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
Holotype- (UMNH VP 28348 in part) (~9 m; subadult or adult) partial
frontals (~145 mm), dorsal rib fragments, mid caudal central fragment,
three fragmentary centra, metacarpal II (51 mm), ilial fragment,
phalanx IV-2 (64 mm), long bone fragments
Referred- ?(NMMNH 50064) tooth fragment (Lewis, 2006)
(WSC 1027) nasals, lacrimal, jugal, ?quadratojugal, quadrate, frontals,
braincase, ?ectopterygoid, dentary, angular, four dorsal ribs,
gastralia, proximal caudal vertebra, scapula, coracoid, pubes
(McDonald, Wolfe and Dooley, 2021)
Diagnosis- (after McDonald,
Wolfe and Dooley, 2021) small, tab-like prefrontonasal process;
prefrontolacrimal process is a rugose prominence separated from
prefrontonasal process by striated notch; prefrontonasal process,
prefrontolacrimal process and notch between them are situated on
mediolaterally-broad, dorsoventrally-thick shelf that roofs dorsal end
of vertical prefrontal contact; medial wall of prefrontal contact does
not merge with ventral surface of prefrontonasal process, such that
ventral surface of prefrontonasal process is free and a deep cleft
separates it from the medial wall of the prefrontal contact.
Other diagnoses- McDonald et
al., (2018) originally diagnosed Dynamoterror
based on two frontal characteristics. Yun (2020) found that the
first (prefrontonasal and prefrontolacrimal processes are in close
proximity, separated only by a shallow notch) was shared with Daspletosaurus sp. frontal SDNHM 32701, but that other Daspletosaurus
individuals (e.g. CMN 8506, RTMP 2001.036.0001) lacked it.
Similarly, Yun found the second character (subrectangular, concave,
laterally projecting posterior part of the postorbital suture,
separated from the anterior part by a deep groove) was shared with the Teratophoneus
holotype BYU 8120/9396 but not referred specimen UMNH VP 16690.
Thus these were both considered characters which vary individually in
tyrannosaurines and are not appropriate for diagnoses. While Yun
used this to declare Dynamoterror
a nomen dubium, McDonald et al. (2021) have since suggested new
autapomorphies that are also seen in their new specimen WSC 1027.
Comments- The holotype was
discovered in August 2012. The supposed pedal phalanx IV-4 is far too small
to be referred to the same individual and may be a phalanx IV-2 based
on its elongation, either ornithomimid or tyrannosauroid. This also
calls into question the referral of most postcranial fragments, which
"were collected as float", although the metacarpal and phalanx IV-2 are
of comprable size.
Lewis (2006) reports "a tyrannosaurid tooth fragment" from the Allison
Member at microvertebrate locality NMMNH L-5636, also reported as
"Tyrannosauridae indet." by Lewis et al. (2007). According to the
NMMNH online catalog, this is NMMNH 50064 found in April 2005.
This may belong to
the co-occuring Dynmamoterror, which it can be compared with once the new specimen WSC 1027 is described that preserved teeth.
McDonald et al. (2018) added Dynamoterror to Carr's tyrannosauroid analysis, ending up in a polytomy with non-alioramin tyrannosaurines and outside Tarbosaurus plus Tyrannosaurus. McDonald et al. (2021) later used their new specimen WSC 1027 to recover Dynamoterror as sister to Teratophoneus, with Lythronax the next closest taxon.
References-
Lewis, 2006. A microvertebrate fauna of the Upper Cretaceous (Late
Santonian-Early Campanian) Menefee Formation, northwestern New Mexico.
Geological Society of America Abstracts with Programs. 38(7), 69.
Lewis, Heckert, Lucas and Williamson, 2007. A diverse new microvertebrate fauna
from the Upper Cretaceous (Late Santonian-Early Campanian) Menefee Formation
of New Mexico. Journal of Vertebrate Paleontology. 27(3), 105A.
McDonald, Wolfe and Dooley Jr, 2018. A new tyrannosaurid (Dinosauria:
Theropoda) from the Upper Cretaceous Menefee Formation of New Mexico.
PeerJ. 6:e5749.
Yun, 2020. A reassessment of the taxonomic validity of Dynamoterror dynastes (Theropoda, Tyrannosauridae). Zoodiversity. 54(3), 259-264.
McDonald, Wolfe and Dooley, 2021. New data on the tyrannosaurid dinosaur Dynamoterror,
including a more complete skeleton, from the Menefee Formation (Middle
Campanian) of New Mexico, USA: Implications for tyrannosaurid evolution
in southern Laramidia. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 181.
Lythronax Loewen, Irmis, Sertich,
Currie and Sampson, 2013
L. argestes Loewen, Irmis, Sertich, Currie and Sampson, 2013
Middle Campanian, Late Cretaceous
Wahweap Formation, Utah, US
Holotype- (UMNH VP 20200) maxilla, nasals, jugal, frontal, quadrate, laterosphenoid,
palatine, dentary, splenial, surangular, prearticular, dorsal rib, chevron,
pubes, tibia, fibula, metatarsal II, metatarsal IV
Diagnosis- (after Loewen et al., 2013) sigmoidal lateral margin of maxilla;
ratio of transverse width of anterior portion of nasal to tranverse width of
middle portion greater than 2.5; prefrontal and postorbital contact surfaces
on frontal nearly in contact, separated only by very narrow, deep dorsoventral
groove; presence of distinct suboccular flange on jugal.
Comments- Loewen et al. (2013) recovered this as the sister taxon to
Tarbosaurus + Zhuchengtyrannus + Tyrannosaurus in their
analysis. In the later analysis incliding Carr's characters and more taxa, Lythronax
was recovered as sister to the Daspletosaurus+Tyrannosaurus clade.
References- Loewen, Sertich, Irmis and Sampson, 2010. Tyrannosaurid evolution
and intracontinental endemism in Laramidia: New evidence from the Campanian
Wahweap Formation of Utah. Journal of Vertebrate Paleontology. Program and Abstracts
2010, 123A.
Loewen, Irmis, Sertich, Currie and Sampson, 2013. Tyrant dinosaur evolution
tracks the rise and fall of Late Cretaceous oceans. PLoS ONE. 8(11), e79420.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
Teratophoneus Carr, Williamson,
Britt and Stadtman, 2011
= "Teratophoneus" Carr, 2005
T. curriei Carr, Williamson, Britt and Stadtman, 2011
= "Teratophoneus curriei" Carr, 2005
Late Campanian, Late Cretaceous
Kaiparowits Formation, Utah, US
Holotype- (BYU 826/9402) maxilla
.... (BYU 8120/9396) lacrimal, partial jugal, frontal, squamosal, quadrates
(198.2, 199.8 mm), basisphenoid, basioccipital, prootic, exoccipital-opisthotic,
partial supraoccipital, articular, third(?) cervical vertebra, partial mid caudal
vertebra (97.1 mm), scapula, coracoid
....(BYU 8120/9397) humerus (241.9 mm), ulna (140.6 mm)
....(BYU 9398) dentary
....(BYU 13719) femur (757 mm)
Referred- (UMNH VP 16690) incomplete skull, partial mandibles, atlas,
seven cervical vertebrae, cervical ribs, eight dorsal vertebrae, fourteen dorsal
ribs, two sacral vertebrae, thirty-four caudal vertebrae, nineteen chevrons,
partial ilia, pubes, ischia, femur, tibia, fibula, pedal phalanx, pedal ungual
(Loewen et al., 2013)
(UMNH VP 16691) jugal (Loewen et al., 2013)
Diagnosis- (after Carr et al., 2011) maxilla with steep anterodorsal
margin; maxillary fenestra situated far posterior to the anterior margin of
antorbital fossa; complete overlap on posterior margin of frontal by parietal; distinct angle in
posterior margin of postorbital process of jugal; basioccipital restricted to
midline of basisphenoid recess as a strut; transversely oriented occiput (where
the paroccipital processes extend nearly directly laterally, instead of posterolaterally);
accessory pneumatic foramen in basisphenoid recess; non-invasive basisphenoid
foramen; subotic recess on basisphenoid; ostium of basisphenoid recess that
opens externally; elevated and pedicle-like joint surface for squamosal on pro�tic.
(after Loewen et al., 2013) midpoint of maxillary fenestra situated posterior
to midpoint of space between anterior edge of antorbital fossa and anterior
edge of antorbital fenestra.
Other diagnoses- McDonald et
al. (2021) found that one of Carr et al.'s (2011) suggested characters
(knob at front of joint surface for quadratojugal on jugal) is shared
with the related Dynamoterror.
Comments- The holotype was discovered in 1981, and the taxon named and
described by Carr in his thesis (Carr, 2005) before being officially described
by Carr et al. (2011). While Stadtman et al. (1999) believed the holotype to
be two individuals, it only consists of one. It is resolved in Carr's analysis
of cranial characters as a basal tyrannosaurine. Carr and Williamson (2010)
include the taxon in their phylogenetic analysis as "new genus from Utah",
where it also resolves as a basal tyrannosaurine. Loewen et al. (2013) described
referred specimens and found the genus to be a tyrannosaurine more derived than
a paraphyletic Daspletosaurus, but less so than Bistahieversor,
Lythronax, Tarbosaurus and Tyrannosaurus. Most recently,
Brusatte and Carr (2016) recovered it as a basal tyrannosaurine sister to Nanuqsaurus
and outside the Daspletosaurus+Tyrannosaurus clade.
References- Stadtman, Chure, Scheetz and Britt, 1999. Fossil vertebrates
from the Kaiparowitz Fm. (Late Cretaceous), Grand Staircase-Escalante Monument
(GRST), Utah. Journal of Vertebrate Paleontology. 19(3), 77A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Wiersma, Loewen and Getty, 2009. Taphonomy of a subadult tyrannosaur from the
Upper Campanian Kaiparowits Formation of Utah. Journal of Vertebrate Paleontology.
29(3), 201A.
Carr and Williamson, 2010. Bistahieversor sealeyi, gen. et sp. nov.,
a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea.
Journal of Vertebrate Paleontology. 30(1), 1-16.
Carr, Williamson, Britt and Stadtman, 2011. Evidence for high taxonomic and
morphologic tyrannosauroid diversity in the Late Cretaceous (Late Campanian)
of the American southwest and a new short-skulled tyrannosaurid from the Kaiparowits
formation of Utah. Naturwissenschaften. 98(3), 241-246.
Loewen, Irmis, Sertich, Currie and Sampson, 2013. Tyrant dinosaur evolution
tracks the rise and fall of Late Cretaceous oceans. PLoS ONE. 8(11), e79420.
Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013. Late Cretaceous theropod
dinosaurs of Southern Utah. In Titus and Loewen (eds.). At the Top of the Grand
Staircase: The Late Cretaceous of southern Utah. Indiana University Press. 504-525.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
McDonald, Wolfe and Dooley, 2021. New data on the tyrannosaurid dinosaur Dynamoterror,
including a more complete skeleton, from the Menefee Formation (Middle
Campanian) of New Mexico, USA: Implications for tyrannosaurid evolution
in southern Laramidia. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 181.
Nanuqsaurus Fiorillo and Tykoski,
2014
N. hoglundi Fiorillo and Tykoski, 2014
Late Maastrichtian, Late Cretaceous
Prince Creek Formation, Alaska, US
Holotype- (DMNH 21461) (skull ~600-700 mm) maxillary fragment, partial frontals,
partial parietals, laterosphenoid, anterior dentary
Diagnosis- (after Fiorillo and Tykoski, 2014) thin, anteriorly forked,
median spur of fused parietals that overlaps and separates frontals within sagittal
crest; frontal with long, anteriorly pointed process separating prefrontal and
lacrimal facets; first two dentary teeth/alveoli much smaller (mesiodistal length)
than dentary teeth/alveoli 3-9 (alveolus 1 <35% of alveolus 3 and <25%
of alveolus 4; alveolus 2 <50% of alveolus 3 and <33% of alveolus 4).
Comments- The holotype was discovered in 2006 and mentioned by Fiorillo
and Tykoski (2013) as being possibly not "referrable to the contemporaneous
Albertosaurus sarcophagus". Fiorillo and Tykoski (2014) described
Nanuqsaurus and added it to both Brusatte's and Loewen's tyrannosauroid
analyses and found it to be sister to the Tarbosaurus+Tyrannosaurus
clade. In the later combination of these analyses by Brusatte and Carr (2016),
Nanuqsaurus is resolved as a basal tyrannosaurine sister to Teratophoneus
in their parsimony analysis, and at this grade outside Daspletosaurus+Tyrannosaurus
in their Bayesian analysis.
References- Fiorillo and Tykoski, 2013. Distribution and polar paleoenvironments
of large theropod skeletal remains from the Prince Creek Formation (Early-Late
Maastrichtian) of Northern Alaska. Journal of Vertebrate Paleontology. Program
and Abstracts 2013, 127.
Fiorillo and Tykoski, 2014. A diminutive new tyrannosaur from the top of the
world. PLoS ONE. 9(3), e91287.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
unnamed clade
Definition- (Daspletosaurus torosus + Tyrannosaurus rex)
= Tyrannosauridae sensu Holtz, 2001
Definition- (Aublysodon mirandus + Tyrannosaurus rex) (modified)
Diagnosis- (after Carr, 2005) maxillary fenestra extends or extends medial
to the anterior margin of the antorbital fossa; anterior process of lacrimal
inflated; medial pneumatic recess pierces anterior lacrimal process; orbitonasal
ridge of lacrimal is positioned close to or reaches the posterior margin of
the bone; lateral bounding ridge of the supratemporal fossa on the squamosal
is divided sagittally; posterior squamosal process inflated; frontolacrimal
contact short in dorsal view.
Comments- This was called Tyrannosaurus by Paul (1988) and contains
all published tyrannosaurine species.
References- Paul, 1988. Predatory Dinosaurs of the World. Simon &
Schuster. 464 pp.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. In Tanke and
Carpenter (eds.). Mesozoic Vertebrate Life. 64-83.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Aublysodontinae Nopcsa, 1928
Definition- (Aublysodon mirandus < Tyrannosaurus rex)
(modified from Holtz, 2001)
Comments- Paul (1988) used this taxon to encompass Aublysodon mirandus,
A. molnari (a juvenile Tyrannosaurus), Shanshanosaurus
(a juvenile Tarbosaurus) and potentially the yet unnamed Archaeornithoides
as well. Holtz (1997, 2001) found Aublysodon (based on A. molnari)
and Alectrosaurus (based partially on IGM 100/50 and 100/51, which are
not Alectrosaurus and are probably juveniles) to clade with OMNH 10131
(a specimen described as Aublysodon but now referred to Bistahieversor).
Currie (2000) assigned both Aublysodon and Alectrosaurus to the
subfamily. Yet the characters used to group these taxa together (unserrated
premaxillary teeth; premaxillary teeth with lingual ridge) are found in all
juvenile tyrannosaurines (Currie, 2003; Carr and Williamson, 2004). Aublysodontinae
is therefore a polyphyletic taxon made of juvenile tyrannosaurids. Holtz's (2001)
phylogenetic definition could include the Daspletosaurus stem if A.
mirandus is in fact a Daspletosaurus specimen. This is based purely
on stratigraphy though, and as the lectotype is indistinguishable from presumedly
Tyrannosaurus juvenile premaxillary teeth, it is inappropriate to use
it to define a clade to the exclusion of Tyrannosaurus. A. mirandus
may even be outside the Daspletosaurus + Tyrannosaurus clade,
or closer to Tyrannosaurus than to Daspletosaurus.
References- Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1,
163-188.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Holtz, 1997. Preliminary phylogenetic analysis of the Tyrannosauridae (Theropoda:
Coelurosauria). Journal of Vertebrate Paleontology. 17(3), 53A.
Currie, 2000. Theropods from the Cretaceous of Mongolia. In Benton, Shishkin,
Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. Cambridge
University Press. 434-455.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. In Tanke and
Carpenter (eds.). Mesozoic Vertebrate Life. 64-83.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous
of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Carr and Williamson, 2004. Diversity of late Maastrichtian Tyrannosauridae (Dinosauria:
Theropoda) from western North America. Zoological Journal of the Linnean Society.
142, 479-523.
Aublysodon Leidy, 1868
A. mirandus Leidy, 1868
= Ornithomimus mirandus (Leidy, 1868) Hay, 1930
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Lectotype- (ANSP 9535; lost) (juvenile) premaxillary tooth
Referred- (AMNH 8514) (juvenile) premaxillary tooth (Sahni, 1972)
(YPM-PU 22252) (juvenile) tooth (Molnar and Carpenter, 1989)
(YPM-PU 23328) (juvenile) tooth (Molnar and Carpenter, 1989)
(YPM-PU 23385) (juvenile) tooth (Molnar and Carpenter, 1989)
(YPM-PU 23389) (juvenile) tooth (Molnar and Carpenter, 1989)
(YPM-PU 23390) (juvenile) tooth (Molnar and Carpenter, 1989)
(YPM-PU 23391) (juvenile) tooth (Molnar and Carpenter, 1989)
(YPM-PU 23387) (juvenile) tooth (Molnar and Carpenter, 1989)
Diagnosis- indeterminate within Tyrannosaurinae.
Comments- Leidy (1856) based Deinodon horridus on fourteen teeth
and tooth fragments discovered in the Judith River Group of Montana. Most were
lateral teeth he regarded as different from Megalosaurus only in their
greater labiolingual thickness, but Leidy placed his species in the new genus
Deinodon because of several other teeth which he felt were distinctive.
These were ANSP 9531, 9533, 9534 and 9535, which can all now be recognized as
tyrannosaurid anterior teeth. Cope (1866) described the teeth of Deinodon
as D-shaped, referencing 9533-9535, to distinguish them from his new taxon Laelaps
(later renamed Dryptosaurus). This makes him first reviser of the genus,
and connected the name Deinodon horridus to the D-shaped teeth in Leidy's
syntype series. Cope considered the lateral teeth to belong to Laelaps.
Leidy (1868) created the new taxon Aublysodon mirandus for ANSP 9533-9535,
intending to retain Deinodon horridus for the lateral teeth (at least
ANSP 9530, 9536 and 9541-9543). Cope's 1866 specification of Deinodon
for the D-shaped teeth has priority though, making Aublysodon mirandus
an objective junior synonym of Deinodon horridus. Marsh (1892) followed
Leidy's (1868) assignment of D-shaped teeth to Aublysodon, and considered
ANSP 9535 to be typical of A. mirandus, while ANSP 9533 and 9534 were
considered examples of another unnamed Aublysodon species. A. mirandus
was notable for its lack of serrations compared to 9533 and 9534. This made
ANSP 9535 the lectotype of Aublysodon, which was formalized by Carpenter
(1982). ANSP 9533 and 9534 are thus implicitly the remaining syntypes of Deinodon
(see entry). Lambe (1902) referred ANSP 9535 to Struthiomimus, while
Osborn (1905) and Lambe (1917) thought it was probably not referrable to Deinodon.
Since Carpenter's (1982) designation of it as the lectotype of Aublysodon,
the latter genus has been most often regarded as a taxon of basal tyrannosauroids
or more recently as an unnatural assemblage of juvenile tyrannosaurid remains.
It is a tyrannosaurid premaxillary tooth, being D shaped and labiolingually
wider than long (by 141% at the base). Both carinae are unserrated and the lingual
face has a broad ridge running vertically. The lack of serrations is also seen
in the premaxillary teeth of juvenile Daspletosaurus and Tyrannosaurus
(LACM 28471), while vertical ridges are present in Gorgosaurus and juvenile
Tyrannosaurus as well. Based on stratigraphy, this is probably a juvenile
Daspletosaurus tooth (Currie, 2005). However, while Daspletosaurus
has been found in the equivalent Oldman, Dinosaur Park and Two Medicine Formations,
it has yet to be reported from the Judith River Formation of Montana. There
is thus no particular species of Daspletosaurus A. mirandus can
be referred to, and as it is indistinguishable from juvenile Tyrannosaurus
teeth, Aublysodon is a nomen dubium within Tyrannosaurinae. For this
reason, it is not a senior synonym of Daspletosaurus.
References- Leidy, 1856. Notices of the remains of extinct reptiles and
fishes, discovered by Dr. F.V. Hayden in the badlands of the Judith River, Nebraska
Territory. Proceedings of the Academy of Natural Sciences of Philadelphia. 8(2),
72-73.
Cope, 1866. [On the remains of a gigantic extinct dinosaur, from the Cretaceous
Green Sand of New Jersey]. Proceedings of the Academy of Natural Sciences of
Philadelphia. 18, 275-279.
Leidy, 1868. Remarks on a jaw fragment of Megalosaurus. Proceedings of
the Academy of Natural Sciences of Philadelphia. 1870, 197-200.
Marsh, 1892. Notes on Mesozoic vertebrate fossils. American Journal of Science.
44, 170-176.
Lambe, 1902. New genera and species from the Belly River Series (Mid-Cretaceous).
Geological Survey of Canada Contributions to Canadian Palaeontology. 3(2), 25-81.
Osborn, 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs.
Bulletin of the American Museum of Natural History. 21, 259-265.
Lambe, 1917. The Cretaceous theropodous dinosaur Gorgosaurus. Geological
Survey of Canada, Memoir. 100, 84 pp.
Hay, 1930. Second bibliography and catalogue of the fossil Vertebrata of North
America. Carnegie Institution of Washington. 390(2), 1074 pp.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin
of the American Museum of Natural History. 147(6), 321-412.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek
formations and a description of a new species of theropod. Contributions to
Geology, University of Wyoming. 20(2), 123-134.
Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian, Montana, U.S.A.)
referred to the genus Aublysodon. Geobios. 22, 445-454.
Currie, 2005. Theropods, including birds. In Currie and Koppelhus (eds.). Dinosaur
Provincial Park, a Spectacular Ancient Ecosystem Revealed. Indiana University
Press. 367-397.
Daspletosaurini Voris, Therrien, Zelenitsky and Brown, 2020
Definition- (Daspletosaurus torosus <- Albertosaurus sarcophagus, Alioramus remotus, Teratophoneus curriei, Tyrannosaurus rex) (Voris, Therrien, Zelenitsky and Brown, 2020)
Diagnosis- (after Voris et al.,
2020) extremely coarse subcutaneous surface of maxilla anteroventral to
antorbital fossa; constricted jugal ramus of maxilla; anteroventral
corner of maxilla tapers in a shallow angle (<65 degrees) as
measured between the alveolar margin of the first two alveoli and the
anterior margin of the subcutaneous surface; at least 14 maxillary
teeth (also in Alioramus);
prefrontal that is broad in dorsal view and strongly dorsomedially
arched in anterior view; dentary chin located ventral to either third
alveolus or third interdental plate.
Comments- This was published in a journal pre-proof posted January 23 2020, but this was electronic
and has no ZooBank entry, 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 June 2020.
Reference- Voris, Therrien,
Zelenitsky and Brown, 2020. A new tyrannosaurine
(Theropoda:Tyrannosauridae) from the Campanian Foremost Formation of
Alberta, Canada, provides insight into the evolution and biogeography
of tyrannosaurids. Cretaceous Research. 110, 104388.
Thanatotheristes Voris, Therrien, Zelenitsky and Brown, 2020
T. degrootorum Voris, Therrien, Zelenitsky and Brown, 2020
= Daspletosaurus degrootorum (Voris, Therrien, Zelenitsky and Brown, 2020) Yun, 2020
Middle Campanian, Late Cretaceous
Foremost Formation of the Judith River Group, Alberta, Canada
Holotype- (RTMP 2010.5.7) (subadult) (skull ~800
mm) partial maxilla, lacrimal fragments, jugal fragment, postorbital
fragments, distal quadrate, fragmentary prefrontal, frontals (one
incomplete, one fragmentary), parietal fragment, partial
laterosphenoid, anterior dentaries, posterior surangular
Paratype- (RTMP 2018.16.1) (subadult) maxillary fragment
Diagnosis- (after Voris et al.,
2020) single row of evenly spaced dorsoventrally oriented ridges on
subcutaneous surface of maxilla ventral and anteroventral to antorbital
fossa; transversely rounded and inflated orbital margin of jugal;
prefrontal with two posteriorly projecting prongs articulating with
frontal on ventral surface of skull roof (long, medial, primary prong
and shorter secondary, lateral prong).
Other diagnoses- Yun (2020) found two proposed frontal autapomorphies of Thanatotheristes suggested by Voris et al. (2020) (sagittal crest on frontal extends anterior to supratemporal ridge as a
broad and rounded ridge; lacrimal contact surface on frontal extends
anteromedially at ~60 degrees relative to interfrontal suture) are also present in Daspletosaurus sp. nov. frontal SDNHM 32701, and that the former is also seen in Ajuga Formation frontal TMM 45905-1.
Comments- The holotype
was discovered in 2010 and the paratype in 2018. It was
described and named by Voris et al. (2020) in a journal pre-proof
posted January 23 2020, but this was electronic
and has a blank space after "ZOOBANK ID:", 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 June 2020. Yun (2020) noted "known material of Thanatotheristes degrootorum are very similar to those of Daspletosaurus
(Voris et al., 2020 [in press]) and the number of currently known
morphological differences between these two genera is actually fewer
than those between two Daspletosaurus species, D. torosus and D. horneri (Carr et al., 2017). Therefore, the genus name "Thanatotheristes" should be considered as a junior subjective synonym of Daspletosaurus." Yet the number of differences is obviously limited by the fragmentary nature of Thanatotheristes compared to D. horneri, and as any such decision is arbitrary, it is not followed here unless it's later adopted by other tyrannosaur workers.
Voris et al. (2020) added the taxon to Carr's tyrannosauroid analysis and found it to be sister to Daspletosaurus torosus plus D. "horneri".
References- Voris, Therrien,
Zelenitsky and Brown, 2020. A new tyrannosaurine
(Theropoda:Tyrannosauridae) from the Campanian Foremost Formation of
Alberta, Canada, provides insight into the evolution and biogeography
of tyrannosaurids. Cretaceous Research. 110, 104388.
Yun, 2020. A subadult frontal of Daspletosaurus torosus
(Theropoda: Tyrannosauridae) from the Late Cretaceous of Alberta,
Canada with implications for tyrannosaurid ontogeny and taxonomy.
Palarch's Journal of Vertebrate Palaeontology. 17(2), 1-13.
Daspletosaurus Russell, 1970
Other definitions- (Daspletosaurus torosus <- Tyrannosaurus rex) (Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017)
Diagnosis-
(after Carr, 2005) hornlet is present on the lateral surface of the
posterior lacrimal process (unknown in Thanatotheristes);
anterodorsal squamosal process stops posterior to level of anterior
margin of laterotemporal fenestra (unknown in Thanatotheristes).
(after Voris et al., 2020) dorsal margin of anterior process of
lacrimal concealed in lateral view (unknown in Thanatotheristes);
cornual process of postorbital approaches laterotemporal fenestra
(unknown in Thanatotheristes); anteroposterior ridge along nasal
process of frontal (unknown in Thanatotheristes); distinct
mediolateral ridge along dorsal margin of epipterygoid fossa of
laterosphenoid; ventral keel on vomer (unknown in Thanatotheristes);
posterior pneumatic foramen of palatine positioned posterior to
anterior margin of vomeroptergoid neck (unknown in Thanatotheristes);
prominent interlocking bony papillae on mandibular symphysis.
Not Daspletosaurus- Langston et al. (1989) referred material from the Aguja Formation of Texas as Daspletosaurus, but that has more recently been identified as a new tyrannosaurine (Lehman and Wick, 2013).
Bell (2007) originally identified Daspletosaurus in
the Horseshoe Canyon Formation based on maxilla RTMP 89.17.53, which is actually
referrable to Albertosaurus (Bell and Currie, 2014).
References- Russell, 1970. Tyrannosaurs from the Late Cretaceous of western
Canada. National Museum of Natural Science Publications in Palaeontology. 1,
1-34.
Langston, Standhardt and Stevens, 1989. Fossil vertebrate
collecting in the Big Bend - history and perspective. In Busbey and Lehman (eds.).
Vertebrate paleontology, biostratigraphy, and depositional environments, Latest
Cretaceous and Tertiary, Big Bend area, Texas. SVP 1989 Guidebook. 11-21.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Bell, 2007. The Danek bonebed: An unusual dinosaur assemblage from the Horseshoe
Canyon Formation, Edmonton, Alberta. Journal of Vertebrate Paleontology. 27(3),
46A.
Lehman and Wick, 2013. Tyrannosauroid dinosaurs from the Aguja
Formation (Upper Cretaceous) of Big Bend National Park, Texas. Earth
and Environmental Science Transactions of the Royal Society of
Edinburgh. 103, 1-15.
Bell and Currie, 2014. Albertosaurus (Dinosauria: Theropoda) material
from an Edmontosaurus bonebed (Horseshoe Canyon Formation) near Edmonton:
Clarification of palaeogeographic distribution. Canadian Journal of Earth Sciences.
51(11), 1052-1057.
Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017. A new tyrannosaur
with evidence for anagenesis and crocodile-like facial sensory system.
Scientific Reports. 7:44942.
D. "horneri" Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017
Late Campanian, Late Cretaceous
Upper Two Medicine Farmation, Montana, US
Material- (AMNH 5477) maxilla (~480 mm), partial postorbital, parietal (Carr and Williamson, 2000; described by Delcourt, 2017)
(MOR 590; intended holotype) (~9 m; adult) incomplete skull (894.5 mm),
mandible, partial forelimb including humerus, incomplete hindlimb
including femur (875 mm), tibia (820 mm), fibula (710 mm), astragalus,
calcaneum, distal tarsal III, distal tarsal IV, metatarsal II (461.8
mm), phalanx II-1 (141.5 mm), phalanx II-2 (99.5 mm), pedal ungual II,
metatarsal III (510.8 mm), phalanx III-1 (129.4 mm), phalanx III-2
(92.3 mm), phalanx III-3 (74.9 mm), pedal ungual III, metatarsal IV
(475.1 mm), phalanx IV-1 (98.5 mm), phalanx IV-2 (~79.1 mm), phalanx
IV-3 (59.2 mm), phalanx IV-4, pedal ungual IV (Varricchio and Currie,
1991; described by Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017)
(MOR 553D.9.19.91) ectopterygoid (Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017)
(MOR 553E.7.6.91.196) ectopterygoid (Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017)
(MOR 553S/7.19.0.97; intended paratype) (juvenile) (skull ~496 mm) dentary (Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017)
(MOR 573) (juvenile) frontal (~71 mm) (Carr and Sedlmayr, 2020)
(MOR 1130; intended paratype) incomplete skull, incomplete mandible,
fourth cervical vertebra (~57.4 mm), seventh cervical vertebra, eighth
cervical vertebra (92.7 mm), three dorsal vertebrae, thirteen caudal
vertebrae (~140.6, 142.1, 150.0, 142.1, 145.9, 124.6, 74.4, 45.2 mm),
partial pelvis, hindlimb including femur, tibia (920 mm), fibula,
calcaneum, distal tarsal III, distal tarsal IV, metatarsal II (480.0
mm), phalanx III-2 (112.7 mm), phalanx III-3 (85.2 mm), metatarsal IV
(530.3 mm), phalanx IV-2 (57.4 mm) (Scherzer, 2008; described by Carr,
Varricchio, Sedlmayr, Roberts and Moore, 2017)
(MOR 3068) partial mandible (Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017)
Diagnosis-
(after Carr et al., 2017) wide dental arcade at front of snout, where
maxillary and dentary tooth rows extend distinctly anteromedially and
the first interdental plate of the maxilla is narrow, which resembles
those of the premaxilla where the tooth row is mediolaterally oriented;
dentary distinctly bowed (convex) laterally (possibly also in Tarbosaurus); promaxillary sinus stopping between alveoli 3 and 4 in subadults and adults, as observed in medial view (also in some Tyrannosaurus);
anterior end of choana on maxilla above alveolus 7; inflated dorsal
surface of lacrimal not reaching medial edge of bone (also in Alioramus);
medial pneumatic recess of lacrimal a tall and narrow slot; concave
upper half of orbital margin of lacrimal; entire circumference of
pneumatic recess of squamosal undercut and clearly defined (also in Tarbosaurus); pneumatic foramen penetrating lateral surface of quadratojugal (also in juvenile Tyrannosaurus); short anterior cervical epipophyses (also in Tarbosaurus).
Other diagnoses- Carr et al.
(2017) proposes several additional diagnostic characters, but in their
supplementary info list multiple other tyrannosaurids which exhibit
them- sinuous anterior edge in dorsal view of frontal dorsotemporal
fossa (also in Gorgosaurus, Lythronax, Raptorex and some Tyrannosaurus); joint surface for squamosal on parietal covers base of posterolateral process (also in Gorgosaurus, Albertosaurus and Tyrannosaurus); crista prootica extends onto prootic (also in Gorgosaurus, Albertosaurus, Tarbosaurus and Tyrannosaurus); shallow notch between the basal tubera (less than 40% total height of basioccipital below occipital condyle) (also in Gorgosaurus and some Tyrannosaurus); humerus ~34% length of femur (also in Gorgosaurus, Albertosaurus, Teratophoneus and possibly D. torosus).
Comments- Varricchio and Currie (1991) announced MOR 590 as "a well preserved tyrannosaurid skull and associated leg are identified as Daspletosaurus"
found "during the last few field seasons." The taxon was noted by
Horner et al. (1992) as "an intermediate tyrannosauirid (represented by
three specimens) [which] has cranial characters that suggest an
evolutionary position between Daspletosaurus from the Judith River Formation and Tyrannosaurus from the Hell Creek Formation." The specimen number MOR 590 was first published in Carr (1999) as Daspletosaurus cf. torosus
for a cast of its maxilla. Only brief details were released
regarding the taxon for the next two and a half decades, though it was
generally recognized as a new species of Daspletosaurus. It was finally described and named Daspletosauruis horneri by Carr et al. (2017). However, this paper has no mention of ZooBank
and as of February 14 2020 Daspletosaurus "horneri" lacks an entry on the ZooBank website. 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"), Daspletosaurus "horneri" Carr et al., 2017 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
MOR 1130 was found in 1992 (Hoepfner, 2017 online), but not excavated
until 2000-2001. It was first mentioned by Scherzer (2008) as Daspletosaurus and was briefly noted in a few papers before being described as a paratype of Daspletosaurus "horneri" by Carr et al. (2017).
Carr and Williamson (2000) first noted AMNH 5477 as a Daspletosaurus maxilla. It was later described by Delcourt (2017) as Daspletosaurus sp., though Carr et al. (2017) listed a postorbital and parietal as also preserved and referred it to D. "horneri."
Horner et al. (1992) thought MOR 590 was transitional between
Daspletosaurus torosus and Tyrannosaurus rex, while Holtz (2001)
recovered it in three possible positions- basal tyrannosaurine, sister to Daspletosaurus
torosus and sister to Tarbosaurus + Tyrannosaurus. Loewen et al. (2013) and the Bayesian
analysis of Brusatte and Carr (2016) recover this as closer to Tyrannosaurus
than Daspletosaurus, but the parsimony analysis of the latter paper places
it as sister to a D. torosus+DPF OTU. Most recently, Carr (2005), Carr and Varricchio (2014) and Carr et al. (2017) recovers it sister to a Daspletosaurus torosus
concept that includes Dinosaur Park Formation specimens. They
note it may be a descendant of the latter but was not ancestral to Tyrannosaurus.
References- Varricchio and Currie, 1991. New theropod finds from the
Two Medicine Formation (Campanian) of Montana. Journal of Vertebrate Paleontology.
12 (3), 59A.
Horner, Varricchio and Goodwin, 1992. Marine transgressions and the evolution
of Cretaceous dinosaurs. Nature. 358, 59-61.
Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria).
Journal of Vertebrate Paleontology. 19(3), 497-520.
Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria:
Coelurosauria) from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New
Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 113-146.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Scherzer, 2008. Taphonomy of the Sun River Bonebed, Late Cretaceous
(Campanian) Two Medicine Formation of Montana. Masters thesis, Montana
State University. 109 pp.
Loewen, Irmis, Sertich, Currie and Sampson, 2013. Tyrant dinosaur evolution
tracks the rise and fall of Late Cretaceous oceans. PLoS ONE. 8(11), e79420.
Carr and Varricchio, 2014. A new species of Daspletosaurus from the Upper
Two Medicine Formation (Late Campanian, Cretaceous) of Montana and evidence
for anagenesis in tyrannosaurine evolution. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 103-104.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017. A new tyrannosaur
with evidence for anagenesis and crocodile-like facial sensory system.
Scientific Reports. 7:44942.
Delcourt, 2017. A subadult maxilla of a Tyrannosauridae from the Two
Medicine formation, Montana, United States. Pap�is Avulsos de Zoologia.
57(9), 113‑118.
Hoepfner, 2017 online. Dinosaur species discovered in Montana provides new information about tyrannosaurs. MSU News Service.
Carr and Sedlmayr, 2020. Juvenile tyrannosaurid frontal from the Two
Medicine Formation (Campanian, Late Cretaceous) shows ontogenetic
recapitulation of phylogenetic character acquisition. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 97.
Daspletosaurus Russell, 1970 sensu Paul, 1988
Diagnosis- (after Carr, 2005) postorbital boss approaches laterotemporal
fenestra.
(after Warshaw and Fowler,
2022) pneumatic inflation of lacrimal reaching medial edge of bone
(allometry, ontogeny, or taphonomy?- Warshaw, 2022, unpublished data
via Warshaw and Fowler, 2022); prefrontal oriented
anteromedially (determined from the angle of the prefrontal articular surface on the lacrimal of the holotype of D. wilsoni, which does not preserve a prefrontal; the prefrontal of D.
"horneri" is oriented mediolaterally); cornual process of
postorbital subdivided into two distinct processes; pneumatic
excavation of the squamosal that does not undercut its anteromedial
margin; quadratojugal lacking a pneumatic foramen in its lateral
surface (individual variation?).
Comments- This clade was recovered by Carr (2005), grouping D. torosus
and an undescribed species from the Dinosaur Park Formation together to the
exclusion of what would be named D. "horneri". On this
website, Paul's (1988) subgenus Daspletosaurus is used as a label for
it.
References- Russell, 1970. Tyrannosaurs from the Late Cretaceous of western
Canada. National Museum of Natural Science Publications in Palaeontology. 1,
1-34.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Warshaw and Fowler, 2022. A transitional species of Daspletosaurus Russell, 1970 from the Judith River Formation of eastern Montana. PeerJ. 10:e14461.
D. (D.) torosus Russell, 1970
= Tyrannosaurus torosus (Russell, 1970) Paul, 1987
Middle Campanian, Late Cretaceous
Oldman Formation, Alberta, Canada
Holotype- (CMN 8506) (9 m, 2.3 tons) (adult) skull (1.04 m), mandible
(1.015, 1.02 m), atlas (40 mm), axis (80 mm), cervical vertebrae 3-10 (780 mm),
dorsal vertebrae 1-13 (1.47 m), dorsal ribs, sacrum (752 mm), caudal vertebrae
1-11, chevrons, scapula (772 mm), coracoid (170 mm), furcula (250 mm), humerus
(357 mm), radius (171 mm), ulna (214 mm), carpal, metacarpal I (60 mm), phalanx
I-1 (133 mm), manual ungual I (155 mm), metacarpal II (120 mm), phalanx II-1
(48 mm), metacarpal III (71 mm), ilium (1.104 m), pubes (935, 902 mm), femur
(1 m)
Referred- ?(RTMP 94.12.602) tooth (Schubert and Ungar, 2005)
(RTMP 97.12.223) maxilla (Schubert and Ungar, 2005)
Diagnosis- (after Warshaw and Fowler, 2022) compared to D. "horneri" and D. wilsoni
- premaxillary tooth row oriented anteromedially such that multiple
teeth are visible in lateral view; antorbital fossa extends anterior to
antorbital fenestra; uninflated anterior lacrimal process; anterior
lacrimal process longer than ventral process; tall lacrimal horn;
dorsal quadrate contact of the quadratojugal not broadly visible in
lateral view.
Comments- Although many other specimens are usually referred to this
species, Currie (2003) noted those from the Dinosaur Park Formation belong
to a then undescribed species (now D. wilsoni),
as do some from the Oldman Formation (Miyashita et al., 2013).
Note the characters listed above from Warshaw and Fowler (2022) are
primitive and in their tree funtion to separate Daspletosaurus torosus from a clade of D. wilsoni, D. "horneri" and other tyrannosaurines. Schubert and Ungar (2005) refer to RTMP 94.12.602 as a tooth,
but this is also the number of a much more complete specimen referred to Gorgosaurus.
It is not clear that Gorgosaurus and Daspletosaurus teeth can
be differentiated in any case.
References- Russell, 1970. Tyrannosaurs from the Late Cretaceous of western
Canada. National Museum of Natural Science Publications in Palaeontology. 1,
1-34.
Paul, 1987. Predation in the meat eating dinosaurs. In Currie and Koster (eds.).
Fourth Symposium on Mesozoic Terrestrial Ecosystems, short papers. 173-178.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous
of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Schubert and Ungar, 2005. Wear facets and enamel spalling in tyrannosaurid dinosaurs.
Acta Palaeontologica Polonica. 50(1), 93-99.
Miyashita, Currie and Paulina-Carabajal, 2013. A new species of Daspletosaurus
(Theropoda: Tyrannosauridae) from the Campanian of southern Alberta represented
by a growth series of well-preserved skulls and skeletons. Journal of Vertebrate
Paleontology. Program and Abstracts 2013, 178.
Warshaw and Fowler, 2022. A transitional species of Daspletosaurus Russell, 1970 from the Judith River Formation of eastern Montana. PeerJ. 10:e14461.
D. (D.) wilsoni Warshaw and Fowler, 2022
Late Campanian, Late Cretaceous
Jack's B2, Judith River Formation, Montana, US
Holotype- (BDM 107) (adult) partial skull (1.05 m), dentary,
splenial, four anterior-mid cervical vertebrae, rib, two sacral neural
spines, caudal vertebrae, chevron, metatarsal I
Late Campanian, Late Cretaceous
Oldman Formation, Alberta, Canada
Referred- (RTMP 2001.36.1) skull, skeleton, skin impressions (Currie,
2003)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
?(AMNH 5438; paratype of Daspletosaurus torosus) (1.52
tons; 17 year old adult) dorsal vertebrae 11-13, sacrum (712 mm), caudal vertebrae
1-2, ilium (1.096 m), pubis, ischium, femur (1 m), tibia (870 mm), metatarsal
II (490 mm) (Russell, 1970)
?(AMNH 5346) maxilla (Russell, 1970)
?(CMN 841) (adult) incomplete postorbital (Carr 1996)
?(CMN 350) hindlimb including femur (930 mm), tibia (870 mm), metatarsus (555
mm) (Russell, 1970)
?(CMN 11594) partial skull (partial maxilla, lacrimal, partial jugal, postorbital,
prefrontals, frontals, parietal, supraoccipital, laterosphenoid, prootic, exoccipital-opisthotic,
basisphenoid, basioccipital), dentaries (Russell, 1970)
?(CMN 11841) frontal, parietal, braincase (Carr, 1996)
(FMNH PR308, originally referred to Gorgosaurus libratus; = AMNH 5336)
(1.79 tons; 21 year old adult) partial skull (980 mm), mandible (~990 mm), skeleton
(femur ~960 mm) (Carr, 1999)
?(NHMUK R4863) premaxilla, maxilla, dentary, hyoid (Russell, 1970)
?(RTMP 82.13.1) (adult) skull (Carr, 1999)
?(RTMP 83.38.1) (adult) skull (Carr, 1999)
(RTMP 85.62.1) (adult) fragmentary skull, fragmentary skeleton including femur
(1 m), tibia (883 mm) and metatarsal III (567.5 mm) (Carrano, 1998)
(RTMP 92.36.1220) skull, skeleton (Carr, 1999)
(RTMP 94.143.1) (Carr, 2005)
?(RTMP 94.218.1) (juvenile) skull (Carr, 1999)
?(RTMP coll.) skull (Currie and Russell, 2005)
?(SDNH 32701) frontal (128 mm) (skull ~753 mm) (Currie, 2003)
?(UA 11) femur (1 m), metatarsal IV(490 mm) (Russell, 1970)
? material (Ryan et al., 2001)
Late Campanian, Late Cretaceous
Upper Two Medicine Farmation, Montana, US
(TA.1997.002.057) (mandible 890 mm) partial dentary (Currie et al., 2005)
....(TA.1997.002.163) metatarsal III (530 mm)
....(TA.1997.002.168) nasals
....(TA.1997.002.264) pedal phalanx
....(TA.1997.002.302) dentary
....(TA.1997.002.385) manual ungual I
....(TA.1997.002.388) lacrimal
....(TA.1997.002.390) surangular (430 mm)
....(TA.1997.002.423) maxilla
....(TA.1997.002.487) maxilla
....(TA.1997.002.496) metatarsal IV(?)
....(TA.1997.002.563) lacrimal
....(TA.1997.002.648) pedal phalanx IV-1
?...(TA.1997.002.781) ilium (1.085 m)
....(TA.1997.002.834) quadrate (232 mm)
....(TA.1997.002.899) quadrate (232 mm)
....(TA.1997.002.1384) jugal
....(TA.1997.002.1435) premaxilla (67 mm)
(TA.1997.002.064) (~7 m) fragmentary premaxilla (60 mm) (Currie et al., 2005)
....(TA.1997.002.071) pedal phalanx
....(TA.1997.002.140) dentary fragment
....(TA.1997.002.200) metatarsal(?) fragment
....(TA.1997.002.316) metatarsal ?IV (458 mm)
?...(TA.1997.002.350) metacarpal II
?...(TA.1997.002.395) manual phalanx II-2(?)
....(TA.1997.002.710) furcula (155 mm)
....(TA.1997.002.1440) ilium (~910 mm)
(TA.1997.002.223) pedal phalanx IV-4(?) (Currie et al., 2005)
?...(TA.1997.002.2) pedal phalanx III-4(?)
....(TA.1997.002.232) distal metatarsal II
....(TA.1997.002.318) pedal phalanx
....(TA.1997.002.321) pedal phalanx III-2
....(TA.1997.002.682) maxilla
....(TA.1997.002.787) distal metatarsal III
....(TA.1997.002.1239) ischium
....(TA.1997.002.1282) dentary
....(TA.1997.002.1308) quadratojugal
....(TA.1997.002.1428) pubis
....(TA.1997.002.1436) maxilla
....(TA.1997.002.1437) ilium (680 mm)
(TA.1997.002.516) pedal phalanx II-1(?) (Currie et al., 2005)
(TA.1997.002.838) pedal ungual IV(?) (Currie et al., 2005)
(TA.1997.002.1383) postorbital (Currie et al., 2005)
(TA.1997.002 coll.) over 1400 elements and fragments including teeth, vertebrae,
ribs (Currie et al., 2005)
Diagnosis- (after Miyashita et al., 2013) dorsal process of premaxilla
extending posteriorly for more than half the diameter of the external naris;
lacrimal that is 1.5 times anteroposteriorly longer than dorsoventrally tall;
pronounced temporal margin of postorbital.
Other diagnoses- while
Miyashita et al. (2013) claimed "a maxillary tooth count greater than
15" distinguished the Dinosaur Park taxon (based on TMP 2001.36.1) from
D. torosus, 15 teeth are present in FMNH PR308 and the D. wilsoni type.
Warshaw and Fowler (2022) use "a rostrocaudally elongate and
dorsoventrally narrow mylohyoid foramen of the splenial" to diagnose D. wilsoni, but this is absent in CMN 11594, FMNH PR308 and RTMP 94.143.1.
Comments- Currie (2003) noted specimens of Daspletosaurus from
the Dinosaur Park Formation appear to represent a distinct species from the
holotype of D. torosus,
citing a paper in preperation by Currie and Bakker. Miyashita et al.
(2013) elaborate using RTMP 2001.36.1 as their example of the new
species. As this is from the Oldman Formation, a simple Oldman
vs. Dinosaur Park separation wouldn't seem to apply and thus the
referral of most Dinosaur Park specimens to this species above should be
considered temporary pending a more detailed review. Loewen et
al. (2013) recover this taxon as just basal to Daspletosaurus+Tyrannosaurus,
but it has not been separated from D. torosus
in most other analyses. Warshaw and Fowler (2022) used BDM 107,
found in 2017 from the Judith River Formation (at a position which
"likely ... corresponds in age to the lower to middle part of the
Dinosaur Park Formation"), as the holotype of a new Daspletosaurus species separate from D. torosus and D. "horneri"- D. wilsoni. They listed characters shared between D. wilsoni and the Dinosaur Park taxon to the exclusion of D. torosus and D. "horneri", and wrote "these features suggests a close affinity between D. wilsoni
and the Dinosaur Park taxon, although this could reflect either
taxonomic synonymity or a genuine sister relationship; this designation
is reserved for future studies centered on the Dinosaur Park taxon
(noted as forthcoming by Currie (2003) and Paulina Carabajal et al.,
2021), which has yet to receive a formal description and may reveal
autapomorphies (or synapomorphies with D. wilsoni) not considered here." This site provisionally accepts the hypothesis the Dinosaur Park taxon is D. wilsoni. Warshaw and Fowler recover D. wilsoni as closer to D. "horneri" and other tyrannosaurines than to D. torosus based on Carr's analysis, proposing an anagenetic series that would lead to wilsoni and "horneri" needing new genus names or a paraphyletic Daspletosaurus.
AMNH 5336 was described by Matthew and Brown (1923) as AMNH 5434, which was
repeated in the literature by Russell and others. It was later moved to the
FMNH as PR308. AMNH 5434 is actually a Gorgosaurus specimen which was
called AMNH 5336 by Matthew and Brown.
SDNHM 32701 is a frontal probably discovered in the 1910s and purchased in 1921 (Yun, 2020b). It was listed as Daspletosaurus sp. by Currie, 2003) and fully described by Yun (2020b) as D. torosus because he assigned both Oldman and Dinosaur Park specimens to that species.
Discovered in 1997 and excavated through at least 2002, the TA.1997.002
specimens are from a single bonebed, representing at least three
individuals (Currie et al., 2005). It's uncertain which individuals
TA.1997.002.516, 838, 1383 or the vertebrae and ribs belong to. Preliminarily described as Daspletosaurus sp., Currie et al. state it "likely represents the same species" as D. "horneri" from the same formation "although further study and description of Daspletosaurus will be necessary before the species can be determined with certainty." Interestingly, now that D. "horneri" has been described, TA.1997.002 instead seems to share characters with D. torosus
plus D. wilsoni. In particular, the
maxillary fenestra is longer than tall, the upper half of the lacrimal
orbital margin is convex, the lacrimal horn is tall, the dorsal margin
of the posterior postorbital process is convex, and the surangular
shelf overhangs the posterior foramen.
References- Matthew and Brown, 1923. Preliminary notices of skeletons
and skulls of Deinodontidae from the Cretaceous of Alberta. American Museum
Novitates. 89, 10 pp.
Russell, 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National
Museum of Natural Science Publications in Palaeontology. 1, 1-34.
Carr, 1996. Tyrannosauridae (Dinosauria: Theropoda) from the Dinosaur Park Formation
(Judith River Group, Upper Cretaceous: Campanian) of Alberta. Masters Thesis.
University of Toronto. 358 pp.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis. The University of Chicago. 424
pp.
Makovicky and Currie, 1998. The presence of a furcula in tyrannosaurid theropods,
and its phylogenetic and functional implications. Journal of Vertebrate Paleontology.
18(1), 143-149.
Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria).
Journal of Vertebrate Paleontology. 19(3), 497-520.
Carr and Williamson, 2001. Resolving tyrannosaurid diversity: Skeletal remains
referred to Aublysodon belong to Tyrannosaurus rex and Daspletosaurus.
Journal of Vertebrate Paleontology. 21(3), 38A.
Ryan, Russell, Eberth and Currie, 2001. The Taphonomy of a Centrosaurus
(Ornithischia: Ceratopsidae) bone bed from the Dinosaur Park Formation (Upper
Campanian), Alberta, Canada, with comments on cranial ontogeny. Palaios. 16,
482-506.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous
of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Currie, Hurum and Sabath, 2003. Skull structure and evolution in tyrannosaurid
dinosaurs. Acta Palaeontologica Polonica. 48(2), 227-234.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Currie, 2005. Theropods, including birds. In Currie and Koppelhus (eds.). Dinosaur
Provincial Park, a spectacular ecosystem revealed. Indiana University Press.
367-397.
Currie and Russell, 2005. The geographic and stratigraphic distribution of articulated
and associated dinosaur remains. In Currie and Koppelhus (eds.). Dinosaur Provincial
Park, a spectacular ecosystem revealed. Indiana University Press. 537-569.
Currie, Trexler, Koppelhus, Wicks and Murphy, 2005. An unusual multi-individiual
tyrannosaurid bonebed in the Two Medicine Formation (Late Cretaceous, Campanian)
of Montana (USA). In Carpenter (ed.). The Carnivorous Dinosaurs. 313-324.
Loewen, Irmis, Sertich, Currie and Sampson, 2013. Tyrant dinosaur evolution
tracks the rise and fall of Late Cretaceous oceans. PLoS ONE. 8(11), e79420.
Miyashita, Currie and Paulina-Carabajal, 2013. A new species of Daspletosaurus
(Theropoda: Tyrannosauridae) from the Campanian of southern Alberta represented
by a growth series of well-preserved skulls and skeletons. Journal of Vertebrate
Paleontology. Program and Abstracts 2013, 178.
Hone and Tanke, 2015. Pre- and postmortem tyrannosaurid bite marks on the remains
of Daspletosaurus (Tyrannosaurinae: Theropoda) from Dinosaur Provincial
Park, Alberta, Canada. PeerJ. 3:e885.
Bell, Campione, Persons, Currie, Larson, Tanke and Bakker, 2017. Tyrannosauroid
integument reveals conflicting patterns of gigantism and feather evolution.
Biology Letters. 13: 20170092.
Yun, 2020a. A reassessment of the taxonomic validity of Dynamoterror dynastes (Theropoda, Tyrannosauridae). Zoodiversity. 54(3), 259-264.
Yun, 2020b. A subadult frontal of Daspletosaurus torosus
(Theropoda: Tyrannosauridae) from the Late Cretaceous of Alberta,
Canada with implications for tyrannosaurid ontogeny and taxonomy.
Palarch's Journal of Vertebrate Palaeontology. 17(2), 1-13.
Warshaw and Fowler, 2022. A transitional species of Daspletosaurus Russell, 1970 from the Judith River Formation of eastern Montana. PeerJ. 10:e14461.
D. (D.) sp. indet. (Carr, 1999)
Middle-Late Campanian, Late Cretaceous
Oldman or Dinosaur Park Formation, Alberta, Canada
Material- (RTMP 80.16.924) frontal, parietal
(RTMP 83.30.1) lacrimal
(RTMP 84.60.1) postorbital
(RTMP 91.36.403) frontal
(RTMP 94.172.115) maxilla
(RTMP 98.48.1) maxilla, nasal
Comments- These elements may belong to either D. torosus or the
undescribed Dinosaur Park species, depending on which formation they were discovered
in (not mentioned by Carr).
Hwang and Claire (2010) mention identifying a tooth UCMP 150589 as Daspletosaurus,
but the UCMP online database lists it as a hadrosaur, so this may be a typo.
References- Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria,
Coelurosauria). Journal of Vertebrate Paleontology. 19(3), 497-520.
Hwang and Claire, 2010. Species and genus-level variation in the tooth enamel
microstructure of tyrannosaurid dinosaurs. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 109A.
D. sp. (Maltese, 2009)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(CMC VP15826; = RMDRC 06-005; Pete III) (11 m) 70%
complete skeleton including jugal, quadratojugals, quadrates,
ectopterygoid, pterygoid, splenial, ?surangular, cervical vertebra,
dorsal vertebrae, dorsal ribs, gastralia, sacrum, first to ninth caudal
vertebrae, tenth and twelfth caudal neural arches, fourteenth caudal
vertebra, sixteenth-twenty-sventh caudal vertebrae, twenty-ninth caudal
vertebra, thirty-first caudal vertebra, thirty-third caudal vertebra,
thirty-fifth caudal vertebra, thirty-seventh-thirty-ninth caudal
vertebrae, twenty-six chevrons, scapulocoracoids (1.1 m), forelimbs
including humeri (~349 mm), radii (~174 mm), ulnae, metacarpal I,
phalanx I-1, metacarpals II (~117 mm), incomplete phalanges II-1,
phalanx II-2, ilia (1.115 m), femur (~1.037 m), tibia, fibula,
astragalus, calcaneum, distal tarsal, metatarsal I, phalanx I-1, pedal
ungual I, metatarsal II (460 mm), phalanx II-1, phalanx II-2, pedal
ungual II, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual
III, metatarsal IV, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal
ungual IV, metatarsal V
Comments- RMDRC 06-005, nicknamed Pete III, was discovered in 2005 (Maltese,
2009). As of 2018 it is on display at the Cincinnati Museum Center
(Maltese, 2018 online). While undescribed, most of the preserved bones
are photographed on the RMDRC blog. Unfortunately, few cranial
elements are preserved and the postcrania of D. "horneri" remain
unillustrated and undescribed aside from measurements. Of listed
differences between Daspletosaurus species, Pete III does have
'anteroventral margin of quadrate's orbital process extends along a
steep 45-degree angle' supposedly shared with "horneri", while torosus
shows a shallow angle. The quadratojugal lacks a lateral pneumatic
foramen unlike adult "horneri" specimen MOR 1130, which Pete III is
larger than (qj height ~229 mm vs. ~199 mm), but the variation of this
feature is unknown, as its apparent absence in the smaller subadult
"horneri" intended holotype (qj height 151.4 mm) could easily be
individual difference instead of ontogenetic difference. The jugal
does lack Thanatotheristes' autapomorphic transversely rounded orbital
margin. It is here referred to Daspletosaurus sp. pending future work.
References- Maltese, 2009. Difficult excavation and preparation of a large Daspletosaurus specimen. Proceedings of the First Annual Fossil Preparation and Collections Symposium. 63-68.
Maltese, 2018 online. Pete III Final Update: In Its Forever Home. RMDRC paleo lab. 12-3-2018.
D? sp. (Carr and Williamson, 2000)
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-22722) partial caudal vertebra
?...(NMMNH P-25083) femur (883 mm)
?...(NMMNH P-27470) anterior dentary, caudal neural arch, caudal centrum, partial
ilium
Comments- This may belong to the same individual and was referred to
cf. Daspletosaurus sp. by Carr and Williamson (2000). However, they also
believed NMMNH P-25049 and OMNH 10131 to be Daspletosaurus, while these
have been referred to Bistahieversor by Carr (2005). The present specimen
may belong to thsat genus as well.
References- Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria:
Coelurosauria) from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New
Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 113-146.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
D? sp. indet. (Demar and Breithaupt, 2006)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- (UW 34823) premaxillary tooth
Reference- Demar and Breithaupt, 2006. The nonmammalian vertebrate microfossil
assemblages of the Mesaverde Formation (Upper Cretaceous, Campanian) of the
Wind River and Bighorn Basin, Wyoming. In Lucas and Sullivan (eds.). Late Cretaceous
Vertebrates from the Western Interior. New Mexico Museum of Natural History
& Science Bulletin. 35, 33-53.
D? sp. indet. (Sullivan, 2006)
Late Campanian, Late Cretaceous
Fossil Forest Member of Fruitland Formation, New Mexico, US
Material- (SMP VP-1658) two teeth
(SMP VP-1693) incomplete pedal phalanx
Comments- No justification for referring these specimens to cf. Daspletosaurus
sp. was given, and it's quite possible they belong to another tyrannosauroid
taxon. Significantly, Sullivan (2006) lists Daspletosaurus as being present
in the Kirtland Formation, based on Carr and Williamson's (2000) identification
of several specimens, most of which have recently been referred to Bistahieversor
(Carr, 2005). It's quite possible the present specimens belong to this
genus as well, though they may be too fragmentary to assign to any
genus.
References- Carr and Williamson, 2000. A review of Tyrannosauridae (Dinosauria:
Coelurosauria) from New Mexico. In Lucas and Heckert (eds.). Dinosaurs of New
Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 113-146.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Sullivan, 2006. Ah-shi-sle-pah Wilderness Study Area (San Juan Basin, New Mexico):
A paleontological (and historical) treasure and resource. New Mexico Museum
of Natural History and Science Bulletin. 34, 169-174.
undescribed possible daspletosaurin (Varricchio, 2001)
Early Campanian, Late Cretaceous
Lower Two Medicine Farmation, Montana, US
Material-
(OTM 200) dentary, splenial, several teeth, cervical vertebrae, dorsal vertebrae,
ribs, sacral vertebrae, caudal vertebrae, chevrons, partial ilia, pubes, ischium
Comments- This was tentatively referred to Daspletosaurus by
Varricchio (2001) based on "the typical tyrannosaurid teeth and
concave ventral margin of the dentary", but remains undescribed.
Dentary concavity is unknown for "Thanatotheristes" and is not
diagnostic for Daspletosaurus in any case (e.g. comparable to Appalachiosaurus).
Reference-
Varricchio, 2001. Gut contents from a Cretaceous tyrannosaurid: Implications
for theropod dinosaur digestive tracts. Journal of Paleontology. 75(2), 401-406.
Tyrannosaurinae sensu Sereno, 1998
Definition- (Tyrannosaurus rex <- Albertosaurus sarcophagus,
Daspletosaurus torosus, Gorgosaurus libratus) (modified)
Diagnosis- (after Carr, 2005) maxillary fenestra extends anteromedial
to the anterior margin of the antorbital fossa; antorbital fossa reaches maxillonasal
suture with elongate contact; accessory pneumatic foramen in anterior lacrimal
process is distal in position; joint surface for the quadratojugal on the jugal
extends anteriorly from the ventral jugal margin; posterodorsal jugal process
extends posterodorsally; lingual bar of the dentary flanks anterior two alveoli;
oval scar of the femur is on the posteromedial edge of the bone; the indentation
of the lateral cnemial process of the tibia is anterior to the midlength of
the process; the posteroventral heel of the calcaneum is short or absent.
Comments- This clade was called Tyrannosaurus by several authors
(Carpenter, 1992; Holtz, 2001; Carr, 2005), but the present consensus (Currie
et al., 2003; Hurum and Sabath, 2003; Holtz, 2004; Brusatte et al., 2010; Loewen
et al., 2013; Brusatte and Carr, 2016) is to retain bataar and rex
in separate genera.
References- Carpenter, 1992. Tyrannosaurids (Dinosauria) of Asia and
North America. In Mateer and Chen (eds.). Aspects of nonmarine Cretaceous geology.
Ocean Press. 250-268.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. In Tanke and
Carpenter (eds.). Mesozoic Vertebrate Life. 64-83.
Currie, Hurum and Sabath, 2003. Skull structure and evolution in tyrannosaurid
dinosaurs. Acta Palaeontologica Polonica. 48(2), 227-234.
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.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis. University of Toronto. 1170 pp.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky
and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms.
Science. 329, 1481-1485.
Loewen, Irmis, Sertich, Currie and Sampson, 2013. Tyrant dinosaur evolution
tracks the rise and fall of Late Cretaceous oceans. PLoS ONE. 8(11), e79420.
unnamed tyrannosaurine (Khozatsky, 1957)
Santonian-Early Campanian, Late Cretaceous
Kara-Cheku, Almaty, Kazakhstan
Material- (IZK 33/MP-61) incomplete dentary
Comments- This specimen was discovered in 1950 and originally referred
to Tyrannosaurus aff. bataar by Khozatsky (1957) and Bazhanov
and Kostenko (1958), and later to Tarbosaurus aff. bataar by Nessov
(1995). Averianov et al. (2012) redescribed it, finding the reduced first dentary
alveolus to place it in the clade of derived tyrannosaurines including Tarbosaurus,
Zhuchengtyrannus and Tyrannosaurus. It differs from these taxa
in lacking a rugose symphysis.
References- Khozatsky, 1957. [To the history of trionychid turtles in
Kazakhstan]. Izvestiya Akademii Nauk Kazakhskoi SSR, Seriya Biologicheskaya.
2, 15-30.
Bazhanov and Kostenko, 1958. [Scheme of stratigraphy of Tertiary deposits of
South-Eastern Kazakhstan and Northern Kirghizia in light of paleontological
data]. Materialy po Istorii Fauny i Flory Kazakhstana. 2, 5-16.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii [Dinosaurs of northern Eurasia: new data about
assemblages, ecology, and paleobiogeography]. Institute for Scientific Research
on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Averianov, Sues and Tleuberdina, 2012. The forgotten dinosaurs of Zhetysu (Eastern
Kazakhstan; Late Cretaceous). Proceedings of the Zoological Institute RAS. 316(2),
139-147.
Tyrannosaurus? zhuchengensis
Hu, Cheng, Pang and Fang, 2001
Campanian, Late Cretaceous
Upper Xingezhuang Formation, Wangshi Series, Shandong, China
Syntypes- (NGMC V1777) metatarsal II (531 mm)
?(NGMC V286) tooth
?(NGMC V288) (juvenile) tooth
?(NGMC V1174) (juvenile) tooth
?(NGMC V1773) tooth
Comments- Originally referred to cf. Tyrannosaurus rex by Hu (1973)
and Dong (1979), this material was named Tyrannosaurus zhuchengensis
by Hu et al. (2001) in their Shantungosaurus monograph. Whether a holotype
was specified is uncertain, as the description has not been translated from
Chinese. There is no evidence the material belongs to one individual or one
taxon, and indeed two teeth are from juveniles unlike the other two and the
metatarsal. Hone et al. (2011) reidentified it as a metatarsal II instead of
metatarsal IV. Hone et al. also state the material is indeterminate, so it cannot
be referred to the sympatric Zhuchengtyrannus and undescribed tyrannosaurid
(ZCDM V0030 and V0032), though it may belong to either. Though Olshevsky (DML,
2002) called it Tarbosaurus zhuchengensis based on geography, this combination
has yet to be published.
References- Hu, 1973. A new hadrosaur from the Cretaceous of Zhucheng,
Shantung. Acta Geologica Sinica. 2, 179-202.
Dong, 1979. Cretaceous dinosaurs of Hunan, China. Mesozoic and Cenozoic Red
Beds of South China. In Institute of Vertebrate Paleontology and Paleoanthropology
and Nanjing Institute of Paleontology (eds.). Selected Papers from the "Cretaceous-Tertiary
Workshop". Science Press. 342-350.
Hu, Cheng, Pang and Fang, 2001. Shantungosaurus giganteus. Geological
Publishing House. 139 pp.
Olshevsky, DML 2002. https://web.archive.org/web/20201113094722/http://dml.cmnh.org/2002Dec/msg00674.html
Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011. A new, large tyrannosaurine
theropod from the Upper Cretaceous of China. Cretaceous Research. 32(4), 495-503.
undescribed tyrannosaurine (Hone, Wang, Sullivan, Zhao, Chen, Li, Ji,
Ji and Xu, 2011)
Campanian, Late Cretaceous
Upper Xingezhuang Formation, Wangshi Series, Shandong, China
Material- (ZCDM V0030) dentary
(ZCDM V0032) maxilla
Comments- Hone et al. (2011) note these bones differ from other tyrannosaurids,
including Zhuchengtyrannus, and will be described in a later paper. Sullivan
et al. (2012) state it differs from Zhuchengtyrannus in having a subcutaneous
flange, lacking a horizontal shelf on the lateral face of the maxillary ascending
process, and the shape and position of the maxillary fenestra being more similar
to Tarbosaurus. Indeed, they stated it "could be referable to T.
bataar despite minor differences from previously described maxillae of that
taxon."
References- Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011.
A new, large tyrannosaurine theropod from the Upper Cretaceous of China. Cretaceous
Research. 32(4), 495-503.
Sullivan, Hone, Rothschild, Wang and Xu, 2012. Tyrannosaurid dinosaurs from
the Upper Cretaceous Wangshi Group of Zhucheng, Shandong Province, China: Coexisting
giant carnivores and a tyrant with a toothache. Journal of Vertebrate Paleontology.
Program and Abstracts 2012, 181-182.
Zhuchengtyrannus Hone, Wang,
Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011
Z. magnus Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu,
2011
Campanian, Late Cretaceous
Upper Xingezhuang Formation, Wangshi Series, Shandong, China
Holotype- (ZCDM V0031) (~12 m; adult) maxilla (640 mm), dentary (760
mm)
Diagnosis- (after Hone et al., 2011) horizontal shelf on lateral surface
of the base of the ascending process; rounded notch in the anterior margin of
the maxillary fenestra.
Comments- While originally placed in a trichotomy with Tarbosaurus
and Tyrannosaurus, this was found to be sister to Tarbosaurus
in Loewen et al.'s (2013) analysis. However, it was recovered as sister to Tarbosaurus+Tyrannosaurus
in Brusatte and Carr's (2016) analysis adding Carr's characters and more taxa.
References- Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011.
A new, large tyrannosaurine theropod from the Upper Cretaceous of China. Cretaceous
Research. 32(4), 495-503.
Loewen, Irmis, Sertich, Currie and Sampson, 2013. Tyrant dinosaur evolution
tracks the rise and fall of Late Cretaceous oceans. PLoS ONE. 8(11), e79420.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of tyrannosauroid
dinosaurs. Scientific Reports. 6, 20252.
Tarbosaurini Olshevsky, 1995
Tarbosaurus Maleev, 1955b
= Shanshanosaurus Dong, 1977
= "Maleevosaurus" Pickering, 1984
= Maleevosaurus Carpenter, 1992
= Jenghizkhan Olshevsky, 1995
T. bataar (Maleev, 1955a) Rozhdestvensky, 1965
= Tyrannosaurus bataar Maleev, 1955a
= Gorgosaurus lancinator Maleev, 1955b
= Gorgosaurus novojilovi Maleev, 1955b
= Tarbosaurus efremovi Maleev, 1955b
= Deinodon novojilovi (Maleev, 1955b) Maleev, 1964
= Deinodon lancinator (Maleev, 1955b) Kuhn, 1965
= Aublysodon lancinator (Maleev, 1955b) Charig in Appleby, Charig, Cox,
Kermack and Tarlo, 1967
= Aublysodon novojilovi (Maleev, 1955b) Charig in Appleby, Charig, Cox,
Kermack and Tarlo, 1967
= Shanshanosaurus huoyanshanensis Dong, 1977
= Tyrannosaurus efremovi (Maleev, 1955b) Rozhdestvensky, 1977
pr= Tyrannosaurus "turpanensis" Zhai, Zhang and Tong, 1978
= Tarbosaurus novojilovi (Maleev, 1955b) Olshevsky, 1978
pr= Tyrannosaurus luanchuanensis Dong, 1979
= "Maleevosaurus" novojilovi (Maleev, 1955b) Pickering, 1984
= Aublysodon huoyanshanensis (Dong, 1977) Paul, 1988
= Albertosaurus novojilovi (Maleev, 1955b) Mader and Bradley, 1989
pr= Tarbosaurus "turpanensis" (Zhai, Zhang and Tong, 1978)
Olshevsky, 1991
pr= Tarbosaurus luanchuanensis (Dong, 1979) Olshevsky, 1991
= Maleevosaurus novojilovi (Maleev, 1955b) Carpenter, 1992
= Jenghizkhan bataar (Maleev, 1955a) Olshevsky, 1995
= Jenghizkhan luanchuanensis (Dong, 1979) Olshevsky, 1995
= Tyrannosaurus novojilovi (Maleev, 1955b) Glut, 1997
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Holotype-
(PIN 551-1) (~12.4 m, ~5 tons) partial skull (~1.35 m), dentary,
posterior cervical vertebrae (c3 65 mm), first dorsal vertebra, second
dorsal vertebra (75 mm), third dorsal vertebra (80 mm), fourth dorsal
vertebra (105 mm) (femur ~1.2 m)
Referred-
(IGM 100/65) partial skull, surangular (Hurum and Sabath, 2003)
(IGM 100F/12) incomplete footprint (Currie, Badamgarav and Koppelhus, 2003)
(IGM 100F/14) incomplete footprint (Currie, Badamgarav and Koppelhus, 2003)
(IGM 107/5; = PJC 2000.9) (juvenile) skull including braincase,
mandibles, ?cervical ribs, four ?dorsal vertebrae, ilium, ischium,
tibia, pedal phalanges (Currie, 2001)
(IGM coll.; 930928 NG WTB) cranial fragment (Watabe and Suzuki, 2000a)
?(IGM coll.; PJC.2001.14) proximal scapula (Currie, 2002)
(IGM coll.) premaxillary tooth (Currie, 2001; Currie, 2003?)
(IGM coll.)
frontal (Currie, 2001)
(IGM coll.)
metatarsal IV (Currie, 2001)
(PIN 551-2; holotype of Tarbosaurus efremovi) (adult) skull,
mandibles, incomplete skeleton including atlas, axis, incomplete
post-axial cervical vertebrae (~c3-c8 80 mm), dorsal vertebrae (mid
dorsal 100 mm), dorsal ribs, (sacrum 700 mm) five sacral vertebrae (170
mm), chevrons, humerus, incomplete ilia (1.080 m), pubes (860 mm),
ischia (750 mm), femora (970 mm), tibia (850 mm), fibula (780 mm),
incomplete astragalus (170 mm trans), calcaneum, metatarsal I (~90 mm),
phalanx I-1 (75 mm), pedal ungual I (55 mm), metatarsal II (455 mm),
phalanx II-1 (140 mm), metatarsal III (540 mm), phalanx III-1 (135 mm),
phalanx III-2 (100 mm), phalanx III-3 (80 mm), pedal ungual III (95
mm), metatarsal IV (510 mm), phalanx IV-1 (100 mm), phalanx IV-3 (60
mm), phalanx IV-4 (45 mm), metatarsal V (~220 mm) (Maleev, 1955b)
(PIN 551-3; paratype of Tarbosaurus efremovi?) (7.7 m, 2.1 tons) skull (~1.130 m), mandible (~1.090 m), skeleton including femur (970 mm) and metatarsus
(546 mm) (Maleev, 1974)
(PIN 551-4; paratype of Tarbosaurus efremovi) incomplete skeleton (Maleev, 1974)
(PIN 551-6) incomplete scapulocoracoid (Maleev, 1974)
(PIN 551-91) partial maxilla (Maleev, 1974)
(PIN 551 coll.; paratypes of Gorgosaurus novojilovi) numerous fragments (Maleev, 1955b)
(PIN 551 coll.) (six individuals) fragmentary skeletons (Hurum and Sabath, 2003)
(ZPAL MgD-I/4; Nemegt, No. 1) partial skull (1.11 m), partial mandible
(dentary 480 mm), presacral fragments, couple dorsal ribs, 13 sacral
and proximal caudal vertebrae, chevrons, forelimb fragments, incomplete
ilium, proximal ischium, femur (970 mm), tibia, fibula, metatarsal I,
phalanx I-1, pedal ungual I, phalanx II-1, phalanx II-2, metatarsal III
(555 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III
(Gradzinsky, 1970)
(ZPAL MgD-I/26) fragmentary maxilla (Hurum and Sabath, 2003)
(ZPAL MgD-I/29; Nemegt No. 2?) partial skull, mandible, six cervical vertebrae, eleven
ribs, five sacral vertebrae, twenty-two caudal vertebrae, humerus,
distal radius, distal ulna, manual digit I, ilium, incomplete pubis,
proximal ischium, femur (580 mm), tibia (590 mm), metatarsus (410 mm),
pes, fragmentary hindlimb (Hurum and Sabath, 2003)
(ZPAL MgD-I/52) dentary tooth (Hurum and Sabath, 2003)
(ZPAL MgD-I/109) (large) skull (Hurum and Sabath, 2003)
(ZPAL MgD-I/178) fragmentary skull, vertebrae, femur (Hurum and Sabath, 2003)
(uncollected?) skeleton (Watabe and Suzuki, 2000a)
(uncollected?) partial skeleton including caudal vertebrae and hindlimbs (Watabe and Suzuki, 2000a)
(destroyed) postcranial skeleton, skin impressions (Watabe and Suzuki, 2000a)
Early Maastrichtian, Late Cretaceous
Altan Uul, Nemegt Formation, Mongolia
(IGM 100/67) fragmentary skull, braincase (Hurum and Sabath, 2003)
(PIN 553-1; holotype of Gorgosaurus lancinator) (~9 m) incomplete skull (970 mm), mandibles (830 mm), four partial
dorsal vertebrae, caudal vertebrae, metacarpal I, metacarpal II, metatarsals, phalanges and unguals
(Maleev, 1955b)
(PIN 553-2) two distal caudal vertebrae (Maleev, 1974)
(PIN 553-3) braincase (Saveliev and Alifanov, 2007)
(ZPAL MgD-I/5) (large) maxilla, quadrate, mandibles (one fragmentary), fragments of eleven ribs,
fragmentary ilia, fragmentary pubis, ischia, hindlimb, metatarsal (Hurum and
Sabath, 2003)
(ZPAL MgD-I/34) cranial fragment, splenial (Hurum and Sabath, 2003)
(ZPAL MgD-I/38) (large) incomplete skull, twelve rib fragments, distal femur,
distal tibia, metatarsal III, metatarsal IV, phalanx IV-1 (Hurum and Sabath,
2003)
(ZPAL MgD-I/44) premaxilla, maxilla, nasal, lacrimal, mandible (Hurum and Sabath,
2003)
(ZPAL MgD-I/45) maxilla, mandible (Hurum and Sabath, 2003)
(ZPAL MgD-I/46) seven cranial fragments, fragmentary mandible, two partial ribs (Hurum
and Sabath, 2003)
(ZPAL MgD-I/67) jugal (Hurum and Sabath, 2003)
(ZPAL MgD-I/93) endocast (Hurum and Sabath, 2003)
(ZPAL MgD-I coll.) (large) skeleton (Kielan-Jaworwska and Barsbold, 1972)
(ZPAL MgD-I coll.) (small) incomplete skeleton (Kielan-Jaworwska and Barsbold, 1972)
?(ZPAL MgD-I coll.) astragalus (105 mm) (Osmolska and Roniewicz, 1970)
(uncollected?) elements (Watabe and Suzuki,
2000c)
(uncollected?) elements (Watabe and Suzuki, 2000c)
(uncollected?) elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
(HMNS coll.) coracoid (Matsumoto, Hashimoto and Sonoda, 2000)
(IGM 100/59) skull (~976 mm), mandible, postcranial skeleton (Barsbold, 1983)
(IGM 100/60) skull, postcranial skeleton (Hurum and Sabath, 2003)
(IGM 100/61) fragmentary skull, postcranial skeleton (Hurum and Sabath, 2003)
(IGM 100/62) fragmentary skull, postcranial skeleton (Hurum and Sabath, 2003)
(IGM 100/70) (medium) fragmentary skull, sclerotic ring, vertebra (Hurum and
Sabath, 2003)
(IGM 107/2) (skull 1.22 m) premaxilla, lacrimal, prefrontal, frontal, parietals,
squamosal, vomer, pterygoid, dentary, surangular, complete skeleton including
femur (1.12 m) and pes (Hurum and Sabath, 2003)
(IGM 107/3) skull (Hurum and Sabath, 2003)
(IGM 107/6A; PJC 2000.25) frontals (81 mm trans interorbital), skull
fragments (uncollected), postcranial fragments (uncollected), skin
impressions (Carpenter, 1997)
(IGM 107/7; 060812 BgT TBM or 060812 BgT-N TBM Tarbo) (2-3 year old juvenile) skull (290 mm), sclerotic
ring, mandibles, posterior dorsal vertebrae, dorsal ribs, eight proximal caudal
vertebrae, proximal chevrons, scapula, coracoid, humerus, radius, ulna, metacarpal
I, phalanx I-1, metacarpal II, phalanx II-1, phalanx II-2, metacarpal III, ilium,
femora (303 mm), tibiae, fibula, astragali, calcaneum, metatarsals II, metatarsals
III, metatarsals IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4,
pedal ungual IV, pedal phalanges, metatarsals V (Watabe, Suzuki, Tsogtbaatar,
Tsubamoto and Saneyoshi, 2010; described by Tsuihiji
et al., 2011)
(IGM coll.; ?=PJC 2000.8) specimen including frontals and occipital condyle (Currie, 2001; Currie, 2003?)
(IGM coll.; 940824 BgT TSGT; = 940823-BgT-TSOGT?) (small) complete
postcranial skeleton including cervical vertebrae, dorsal vertebrae,
pelvis, femora, fibula, metatarsal, digits (Watabe and Suzuki, 2000b)
(IGM coll.; 940826 BgT OTGN) mandible, postcrania (Watabe and Suzuki, 2000b)
(IGM coll.; 950622 BgT Tarbo. A) caudal vertebrae (Suzuki and Watabe, 2000a)
....(950622-25 BgT Tarbo. PJ1-9) elements (Suzuki and Watabe, 2000a)
........(950622 BgT Tarbo PJ-1) pes (Matsumoto,
Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi,
2010)
........(950625 BgT Tarbo Pj-8) femur (Matsumoto,
Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi,
2010)
........(950625 BgT Tarbo PJ-9) tibia, fibula, metatarsals phalanges (Matsumoto,
Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi,
2010)
....(950624 BgT Tarbo. B) skull (Suzuki and Watabe, 2000a)
....(950626 BgT Tarbo. C) pelvis including pubis, femur (Suzuki and Watabe, 2000a)
....(950817 BgT Tarbo. D) ribs (Suzuki and Watabe, 2000a)
....(950817 BgT Tarbo. E) gastralia (Suzuki and Watabe, 2000a)
....(950817 BgT Tarbo. F) gastralia (Suzuki and Watabe, 2000a)
(IGM coll.; 980803 BgT NAR) (small) partial skeleton (Suzuki and Watabe, 2000b)
(IGM coll.; 060816 BgT TUI) forelimb (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
(IGM coll.; 060817 BgT MB) ribs, pelvis, metatarsus, pedal phalanges
(Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
(IGM coll.)
tooth (Currie, 2001)
(uncollected?) skeleton (Watabe and Suzuki, 2000a)
(uncollected?) specimen (Watabe and Suzuki, 2000a)
(uncollected?) cranial fragments (Watabe and Suzuki, 2000b)
(uncollected?) skull (Watabe and Suzuki, 2000b)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(IGM coll.; 980808 GT ULZ Tarbo) (small) partial skeleton including
incomplete caudal series, chevrons, pelvis including ischium and
hindlimbs including tibiae, metatarsals, pedal phalanges and pedal
unguals (Suzuki and Watabe, 2000b)
(uncollected?) cranial fragments (Watabe and Suzuki, 2000b)
(uncollected) (large) skeleton (Suzuki and Watabe, 2000b)
(uncollected?) elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
White Beds of Khermeen Tsav, Nemegt Formation, Mongolia
(IGM 100/69) occiput (Hurum and Sabath, 2003)
(IGM coll.; 970716-18 KmT) (at least 2 individuals) skull including
maxillae, jugal, braincase, mandible, seven mid caudal vertebrae, mid
chevron, humerus, pelvis, femur, other elements including metatarsal I,
phalanx I-1, pedal ungual I, metatarsal II (~513 mm), phalanx II-1,
phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx
III-2, phalanx III-3, pedal ungual III, metatarsal IV (~521 mm),
phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual
IV, metatarsal V (Watabe and Suzuki, 2000c)
Early Maastrichtian, Late Cretaceous
Tsagan Khushuu (= Tsagan Ula), Nemegt Formation, Mongolia
(IGM coll.; PIN 552-1; paratype of Tarbosaurus efremovi)
incomplete skeleton missing distal caudal vertebrae including
gastralia, scapula (750 mm), coracoid, humerus (255 mm), radius (110
mm), ulna (115 mm), intermedium, metacarpal I (38 mm), phalanx I-1 (65
mm), manual ungual I, metacarpal II (60 mm), phalanx II-1 (35 mm),
phalanx II-2 (55 mm), manual ungual II, metacarpal III, femur (970 mm),
tibia (870 mm), distal tarsal III, distal tarsal IV, metatarsal II (455
mm), phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (540
mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III,
metatarsal IV (510 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal ungual IV, metatarsal V (Maleev, 1974)
(PIN 552-2; holotype of Gorgosaurus novojilovi) (6.18 m,
juvenile) partial skull (713 mm), dentary (380 mm), (axis to c10 450
mm) cervical vertebrae 3-10, cervical ribs, dorsal vertebrae 1-13,
dorsal ribs, gastralia, sacrum (450 mm), caudal vertebrae 1-45 (c1-24
~1.970 m), chevrons, incomplete scapula (330 mm), coracoid, humerus
(143 mm), radius (108 mm), ulna, manus including metacarpal I (35 mm),
metacarpal II (45 mm) and phalanx II-1 (45 mm), ilium (625 mm), pubis
(460 mm), ischium (390 mm), femur (560 mm), tibia (585 mm), fibula (525
mm), astragalus (125 mm trans), calcaneum, distal tarsal III,
metatarsal II (365 mm), phalanx II-1, phalanx II-2, pedal ungual II,
metatarsal III (420 mm), phalanx III-1 (90 mm), phalanx III-2 (70 mm),
phalanx III-3 (55 mm), pedal ungual III, metatarsal IV (395 mm),
phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
(Maleev, 1955b)
(PIN 552 coll.; paratypes of Tarbosaurus efremovi) many elements (Maleev, 1955b)
(ZPAL MgD-I/3; Tsagan Khushu, No. 2) (5.8 m, 760 kg) incomplete skull
(745 mm), posterior mandibles, cervical series, cervical ribs, dorsal
series, dorsal ribs, gastralia, ten proximal caudal vertebrae, proximal
chevrons, scapulae, coracoids, forelimbs (one fragmentary) including
humerus, radius, ulna, metacarpal I, phalanx I-1, manual ungual I,
metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, ilium,
incomplete pubis, ischium, femora (700 mm), tibiae (700 mm), fibulae,
metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal
III (445 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual
III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx
IV-4, pedal ungual IV, metatarsal V? (Hurum and Sabath, 2003)
(ZPAL MgD-I/31) posterior mandible (Hurum and Sabath, 2003)
(ZPAL MgD-I coll; Tsagan Khushu, No. 4) incomplete skeleton (Gradzinsky, 1970)
(uncollected?) maxilla (Watabe and Suzuki, 2000a)
(uncollected?) elements (Watabe and Suzuki,
2000c)
(uncollected?) elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Ulaan Khushuu, Nemegt Formation, Mongolia
(uncollected?) elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Late Cretaceous?
Nemegt Formation?, Mongolia
Referred-
(IGM 100/66) (juvenile) specimen including nasals and lacrimal (Currie, 2003)
(IGM 100/177) (juvenile) specimen including maxilla (Currie and Dong, 2001)
(IGM 100/777) (juvenile) specimen including premaxillae, maxilla,
nasal, vomer (Currie, 2001)
(IGM 107/1) incomplete skull (991 mm), dentary, coronoid, splenial (Hurum and Sabath, 2003)
(IGM 107/14) (three individuals; subadults) cranial elements including nasals
(293 mm), lacrimal (172 mm), postorbital, squamosal, braincase, postcranial
elements (Tsuihiji, 2010)
(IGM coll.; PJC 2005.07) specimen including braincase (Paulina Carabajal, 2015)
(IGM coll.; PJC 2005.17) specimen including braincase (Paulina Carabajal, 2015)
(IGM coll.; GIN Jap-Mong of Currie, 2003) frontals, parietals (Currie, 2003)
(Tokyo Natural Science Museum coll.) skull, incomplete skeleton (www.paleofile.com)
partial skeleton(s) (Lee, Barsbold, Jacobs and Currie, 2008)
Middle Campanian-Early Maastrichtian, Late Cretaceous
Nemegt or Baruungoyot Formation, Mongolia
(ZPAL MgD-I/16) (ZPAL online)
(ZPAL MgD-I/19) (ZPAL online)
(ZPAL MgD-I/21) (ZPAL online)
(ZPAL MgD-I/28) (ZPAL online)
(ZPAL MgD-I/30) tibia (825 mm), metatarsus (525 mm) (Holtz, 1994)
(ZPAL MgD-I/33) (ZPAL online)
(ZPAL MgD-I/36) (ZPAL online)
(ZPAL MgD-I/54) (ZPAL online)
(ZPAL MgD-I/59) (ZPAL online)
(ZPAL MgD-I/60) (ZPAL online)
(ZPAL MgD-I/61) (ZPAL online)
(ZPAL MgD-I/71) (ZPAL online)
(ZPAL MgD-I/72) (ZPAL online)
(ZPAL MgD-I/76) (ZPAL online)
(ZPAL MgD-I/81) (ZPAL online)
(ZPAL MgD-I/90) (ZPAL online)
(ZPAL MgD-I/175) fragmentary skull (Brusatte, Carr, Erickson, Bever and Norell,
2009)
(ZPAL MgD-I/176) (ZPAL online)
(ZPAL MgD-I/177) (ZPAL online)
Maastrichtian, Late Cretaceous
Subashi Formation, Xinjiang, China
(IVPP V4878; holotype of Shanshanosaurus huoyanshanensis) (2.3 m, 27
kg, juvenile) (skull ~288 mm) premaxilla (lost), maxilla (180 mm), mandible,
tooth (14.4 mm), atlantal centrum, axis (22.3 mm), nine incomplete cervical
vertebrae (anterior cervical 21.2 mm), cervical postzygapophysis, anterior cervical
rib, thirteen incomplete dorsal vertebrae (posterior dorsal 38.7 mm), several
dorsal ribs, scapula (138 mm), coracoid, humerus (88.8 mm), distal pubes, femur
(279 mm), proximal tibiae (Dong, 1977)
?(IVPP coll.; material of Tyrannosaurus "turpanensis") five
teeth, three posterior sacral vertebrae, ilium (Dong, 1977)
Campanian-Early Maastrichtian, Late Cretaceous
Quiba Formation, Henan, China
?(IVPP V4733; holotype of Tyrannosaurus luanchuanensis) five teeth (35.1-36.9
mm), partial vertebra (Dong, 1979)
Diagnosis- (after Carr, 2005) subcutaneous flange extends dorsally from
the main body of the maxilla to block the antorbital fossa from lateral view
(variably present; also in some Alioramus); vertical ridge reinforces
the concave proximal joint surface of pedal phalanx II-2; the medial margin
of the proximal joint surface of pedal phalanx IV-1 is concave.
Original specimens- The holotype was discovered in 1946 (Maleev, 1955a), while the Tarbosaurus efremovi
holotype PIN 551-2 and paratypes (three more "almost complete skeletons
and many miscellaneous bones") were found in 1948-1949 (Maleev,
1955b). Maleev (1974) specifies two of the paratype skeletons are
551-4 and 552-1, while the other is probably PIN 551-3, which is
figured as T. efremovi. The types of Gorgosaurus lancinator (PIN 553-1) and G. novojilovi
(PIN 552-2) were also found in 1948-1949. Maleev died in 1966
with a large tyrannosaurid monograph in preparation which was published
posthumously with editing by Kurzanov and Rozhdestvensky (Maleev,
1974). PIN 552-3 is listed by Ford (paleofile.com) as "incomplete
skull and partial skeleton" and is a typo for PIN 552-2 in Currie
(2003), being described as the "holotype of "Maleevosaurus novojilovi,"
after Maleev 1974." Similarly, PIN 552-4 is listed by Ford as
"partial skeletons" and PIN 555-5 as "partial skeleton", but there is
no evidence in the literature for the existance of these.
Osmolska and Roniewicz (1970) state that associated with the Deinocheirus
holotype found in 1965, "a nearly complete astragalus of carnosaurian
type, 105 mm long (tr.) was found. This seems too small to be assigned
to this specimen, however, and most probably belongs to a
tarbosaur." Barsbold (1983) lists "complete skull and well
preserved postcranial skeleton of one specimen (no. 100/59)" and states
Tarbosaurus has a distribution
of Bugin Tsav. He illustrates the skull and mandible. Hurum
and Sabath (2003) studied numerous ZPAL specimens discovered between
1964 and 1970, and several IGM specimens discovered from 1964-1986 (see
paper for details). They note PIN 552-1 had been transferred to
the IGM, and also that "PIN collections also contains several other
skeletons, including fragmentary remains of at least six individuals
from Nemegt, collected during the Soviet expeditions in the years
1946-1949." Gradzinsky (1970) illustrates ZPAL MgD-I/3 as "Tsagan
Khushu, No. 2" in figure 28, and ZPAL MgD-I/4 as "Nemegt, No. 1" in
figure 29. This leaves ZPAL MgD-I/29 as probably " Nemegt, No.
2", but which specimen is "Tsagan Khushu, No. 4" is uncertain, though
it is probably one of those listed on the ZPAL website.
Kielan-Jaworwska and Barsbold (1972) note that in Summer 1970 "two
incomplete skeletons of Tarbosaurus
sp. and several fragments of the skeletons of the same species" were
found at Nemegt, which are probably ZPAL MgD-I/4 and I/29. They
also say in August or September of 1970 "an incomplete skeleton of Tarbosaurus bataar"
was found at Altan Uul, which is probably ZPAL MgD-I/1-5. The
large and small specimens found in 1971 in Altan Uul III are probably
among those listed on the ZPAL website. Gradzinski et al. (1977)
listed Tarbosaurus bataar as
being known from Bugin Tsav, perhaps based on IGM 100/60-62, found
between 1964 and 1966. They also listed it as being present in
the White Beds of Khermeen Tsav, perhaps based on IGM 100/69 found in
1973. Perle et al. (1994) report "in the immediate vicinity of
the holotype skeleton [of Mononykus; IGM 107/6] the remains of chelonians and the tyrannosaurid Tarbosaurus bataar
were recovered" at Bugin Tsav, which is probably IGM 107/3.
Suzuki and Watabe (2000b) state that in August 1998 they "exposed the
large skeleton of Tarbosaurus discovered in 1987 by Mongolian expedition party" at Gurilin Tsav.
New specimens- Watabe and
Suzuki (2000a) reported a "Tarbosaurus skeleton" found at Bugin Tsav on
September 6 1993. Another Tarbosaurus was found there between
September 14-17 1993. On September 18 1993 a maxilla was
discovered at Tsagan Khushu, then "Tarbosaurus skeletal elements" at
Altan Ula-II on September 19. At Nemegt, they reported a
"Tarbosaurus skeleton (badly weathered)" was found at Central Sayr in
September 24-29, while in the same period a "Tarbosaurus partial
skeleton with hindlimbs - caudals articulated was found at the site
near the Reconnaissance Hill that had been named by the Polish team in
1964." A "Tarbosaurus skull fragment" catalogued as field number
930928 might be part of the latter specimen, as it was from the Western
Sayr so could not belong to the weathered skeleton. Watabe and
Suzuki (2000b) reported a "complete skeleton of a small individual of
Tarbosaurus" (probably field number 940824 BgT TSGT listed as
"Tarbosaurus whole skeleton without skull"), "other skeletons of large
Tarbosaurus and Saurolophus from a single site" (probably "3 incomplete
skeletons of Tarbosaurus" mentioned later) and "fragments of a skull of
Tarbosaurus" found between August 19-25 1994, "a badly preserved
Tarbosaurus skull" found at Bugin Tsav II on August 21, and a
"Tarbosaurus in northeastern site" on August 26 (probably field number
940826 BgT OTGN listed as "Tarbosaurus lower jaw and
postcranials"). They also reported "Tarbosaurus skull fragments"
found on August 28 at Gurilin Tsav. Tsogtbaatar (2004) listed
940823-BgT-TSOGT as a "Skeleton of Tarbosaurus sp.", "incomplete
articulated skeleton of a small tarbosaur" prepared December 1995 to
June 1996 that is probably 940824 BgT TSGT. Suzuki and Watabe
(2000a) reported the specimens found by Saurolophus in 1994 were
collected on June 16-27 1995, as field numbers 950622 BgT Tarbo. A
"Tarbosaurus caudal vertebrae", 950622-25 BgT Tarbo. PJ1-9 "Tarbosaurus
bones", 950624 BgT Tarbo. B "Tarbosaurus skull" and 950626 BgT Tarbo. C
"Tarbosaurus pelvis." They collected the rest of the material on
the way back in August 17 as field numbers 950817 BgT Tarbo. D
"Tarbosaurus ribs", 950817BgT Tarbo. E and 950817 BgT Tarbo. F, both
"Tarbosaurus gastralia." While the 1994 summary states three
individuals were present, the 1995 report states "a skeleton of
Tarbosaurus" was excavated. The pubis was prepared in 1998 and
stored at the HMNS (Matsumoto et al., 2000). Watabe and Suzuki
(2000c) reported "isolated skeletons of Tarbosaurus, including at least
two individuals with similar size" were excavated in mid July 1997 from
the Upper White Beds of Kermeen Tsav, corresponding to field number
970716-18 KmT "Tarbosaurus skull, pelvis, femur, humerus others."
An in situ photo is shown labeled "Skeletons of Tarbosaurus from the
Upper White Bed in Khermeen Tsav. This is also probably the
complete pes photographed by Matsumoto et al. (2010) labeled 97-21-47
(figure 2a). More than 2 individuals are buried at the
site." It shows a maxilla, mid caudals and a chevron.
Tsogtbaatar (2004) listed the specimen as field number 970718 KmT
Tarbo, including"right and left maxillae, brain case, jugal, lower jaw,
femur." They also reported "isolated bones of Tarbosaurus" from
Altan Ula-IV and Tsagan Khushu found on July 21-28 and 25-26 1997
respectively. Isolated bones were also reported from Altan
Ula-III. Suzuki and Watabe (2000b) reported a "partial skeleton
of Tarbosaurus" found on July 30 1998 at Bugin Tsav, given field number
980803 BgT NAR. They also reported "an articulated pelvic-caudal
skeleton of small Tarbosaurus" found at Gurilin Tsav on August 8 1998,
later cited as a "partial skeleton (caudals, pelvic part, and
hindlimbs) of small-sized Tarbosaurus." This was given field
number 980808 GT ULZ Tarbo and photographed in situ as "Tarbosaurus
hind legs and caudal vertebrae at Gurilin Tsav". Matsumoto et al.
(2000) listed an isolated coracoid from Bugin Tsav as being prepared in
early August 1997 and held at the HMNS. Watabe et al. (2010)
reported a number of additional specimens found in 2006. One of
these is field number 060812 BgT TBM (or 060812 BgT-N TBM Tarbo) discovered at Bugin Tsav in 2006,
photographed by Matsumoto et al. (2010) as 2006-04-001 (figure 2b) and
briefly described by Tsuihiji et al. (2007). Its skull was later
described in detail by Tsuihiji et al.
(2011) as IGM 107/7.
Currie (2001) figures a Tarbosaurus snout recovered in 1999, which
matches one of two maxillae illustrated by Currie and Dong (2001) as
"Maxillae of young Tarbosaurus (GIN 100/177)" and that in Currie (2003)
Figure 3F labeled "Young Tarbosaurus bataar, GIN 100/777." Thus
it is assumed to be 100/777 here while the other more slender maxilla
in Currie and Dong's figure is assigned to 100/177. Currie (2001)
announced several new specimens collected in September 2000 from the
Nemegt locality (all sent to the IGM)- a frontal, metatarsal IV,
premaxillary tooth, and a juvenile specimen whose pedal phalanges were
collected, but the rest left to be excavated in 2001 (listed as
PJC.2000.9 by Currie, 2002). This was later catalogued as IGM
107/5, associated with hadrosaurid footprints and figured schematically
by Currie et al. (2003). may be the specimen listed with field
number PJC 2000.26 by Currie (2003), with listed frontal measurements
slightly larger than IGM 107/7. Two footprints found at the same
locality (IGM 100F/12 and 100F/14) were also described by Currie et al.
and referred to Tarbosaurus bataar. They can now be confidently
assigned to Tyrannosauridae instead of Deinocheirus because the latter
has blunt pedal unguals, and are probably Tarbosaurus as even large
Alioramus (e.g. Qianzhousaurus type) are smaller (skull of IGM 100F/12
~1.33 m based on third digit length). IGM 100F/12 preserves scale
impressions. Paulina Carabajal (2015) lists three Tarbosaurus
baatar neurocrania- PJC 2005.07, PJC 2005.17 and PJC 2000.9, all field
numbers that would eventually be given IGM numbers. The "frontals
plus occipital condyle" noted as collected in September 2000 by Currie
(2001) from Bugin Tsav are possibly PJC.2000.8 listed by Currie
(2003). Finally, a tooth was reported by Currie (2001) from Bugin
Tsav. Currie (2002) reported a cf. Tarbosaurus glenoid portion of
a scapula found at the Nemegt locality (PJC.2001.14) found in September
2001. Lee et al. (2008) reported that Tarbosaurus partial
skeletons were discovered by the Korea-Mongolia International Dinosaur
Project in 2006 and/or 2007 from one or more of several Nemegt
Formation localities ("Ulan Khushu, Altan Uul, Bugin Tsav, and Guriliin
Tsav") which have previously known specimens.
A furcula is known
(Sabath pers. comm. to Carpenter and Smith, 2001), but the specimen it
belongs to is unreported. Ford (paleofile.com) lists "Tokyo
Natural Science Museum: Skull and nearly complete skeleton" under
Tarbosaurus efremovi, but further information could not be found.
Skin impressions- Carpenter (1997) stated "impressions of skin around a badly weathered skull of Tyrannosaurus (= Tarbosaurus) bataar
in Mongolia showed the presence of a wattle or bag of skin under the
jaws (Mikhailov, personal communication)" and later (Carpenter, 1999)
said "Tyrannosaurus
did,
however, apparently have either a pelican-like pouch or dewlap based on
an impression of the skin found below a skull in Mongolia (Mikhailov,
personal communication)." Mikhailov (pers. comm. 10-8-2019)
states that the specimen was from the Nemegt Formation and found by
Kurzanov but not collected, being on a "large and heavy stone
plate." He furthermore said the identification as a particular
integumentary structure was Kurzanov's interpretation. This is
possibly the "frontals plus skin impression" reported from Bugin Tsav
by Currie (2001) who stated it was "from one of the skeletons
apparently destroyed by locals looking for material to sell on the
black market" and that the "specimen [was] known for many years."
Currie et al. (2003) and Bell et al. (2017) briefly described and
figured this speciemen as IGM 107/6A. They state the originally
compelete skeleton was poached, leaving only uncollected cranial and
postcranial fragments besides the frontals and two skin patches, which
have an uncertain placement on the body due to the poaching
damage. Currie et al. (2003) states the frontal interorbital
width is 81 mm, which matches field number PJC 2000.25 as listed by
Currie (2003).
Watabe and Suzuki (2000a) reported that in late September 1993 "it
became clear that in the Western Sayr (named by the Polish expedition
team), the articulated postcranial skeleton of Tarbosaurus with skin
impression that had been discovered and left in field in 1992 had been
destroyed by someone." While the timeline would work out with
Carpenter's specimen, it being a posrcranial skeleton would not.
Similarly, it being at Nemegt rather than Bugin Tsav shows this was not
what would become IGM 107/6A.
One or more species?- The holotype specimen was first named Tyrannosaurus
bataar by Maleev (1955a), with smaller specimens subsequently named Tarbosaurus
efremovi, Gorgosaurus lancinator and Gorgosaurus novojilovi
(Maleev, 1955b). Maleev (1964) later transferred the latter two species to Deinodon.
Rozhdestvensky (1965) synonymized all four species into Tarbosaurus bataar,
while Maleev (1974) and Barsbold (1983) used the name Tarbosaurus efremovi
instead. Paul (1988) placed all Nemegt tyrannosaurs into Tyrannosaurus bataar.
These authors all viewed the various sizes and morphologies as a growth series
of one species. Carpenter (1992) separated G. novojilovi as the new genus
Maleevosaurus based on the laterally obsured promaxillary fenestra; small
maxillary fenestra; large, elongate antorbital fenestra; low and slender maxilla;
moderately developed lacrimal horn lacking rugosity; slender jugal; non-rugose
postorbital; slender dentary; tall cervical neural spines; reduced acromion
on scapula; pronounced spur-like obturator process; downcurved ischium; and
metatarsals III and IV don't overlap the metatarsals medial to them much. Carr
(1998; 1999; 2005) has shown the cranial characters are due to ontogeny, while
the only ontogenetic studies of tyrannosaur postcrania that have been published
have dealt with proportions. Nor has individual variation in postcrania been
studied much, though Tyrannosaurus does vary in obturator process size
and ischial curvature. Thus the postcranial characters are here seen as ontogenetic
or individual variation, perhaps even involving preservational effects. Even
Olshevsky currently believes Maleevosaurus to be a juvenile tarbosaur.
Olshevsky (1995) separated Tyrannosaurus bataar from Tarbosaurus efremovi,
placing the former species in the new genus Jenghizkhan because he did
not believe it to be closer to Tyrannosaurus than to Tarbosaurus.
He diagnosed this taxon using a number of seemingly ontogenetic characters-
large size; massive and rugose preorbital and postorbital bars; lacrimal-postorbital
contact; well developed anterior dorsal parapophyses; as well as a couple postcranial
characters of uncertain significance- tall anterior dorsal neural arches; well
developed anterior dorsal neural arch laminae. Olshevsky claimed since the Gorgosaurus
lancinator holotype (PIN 553-1) is a smaller specimen than the Tarbosaurus
efremovi holotype (PIN 551-2), yet shows the cranial characters of PIN 551-1,
it is a juvenile Jenghizkhan and the characters are not ontogenetic.
Rugosity can be individually variable as well as ontogenetically variable. In
Tyrannosaurus, FMNH PR2081 has more young features than its size suggests
it should (Carr, 2005), and this may be true for the holotype of Tarbosaurus
efremovi as well. Vertebral characters have not been examined for taxonomic,
ontogenetic or individual variation in any tyrannosaurids, so their significance
in diagnosing Jenghizkhan is unclear. Although variation in Nemegt tyrannosaurines
hasn't been studied in depth, basically all researchers find no justification
for recognizing more than one species - Tarbosaurus bataar (Currie, 2003;
Hurum and Sabath, 2003; Holtz, 2004; Carr, 2005).
Shanshanosaurus- Discovered in 1964-1966, Shanshanosaurus
was described from the Subashi Formation of Xinjiang, China (Dong, 1977). Dong
placed it its own family, Shanshanosauridae, close to the Tyrannosauridae within
Carnosauria. He posed but dismissed the possibility it was a juvenile tyrannosaurid
based on his incorrect interpretation of the odontoid process being fused to
the axis and vague cranial and mandibular characters. Paul (1988) thought Shanshanosaurus
was related to Aublysodon mirandus and LACM 28471 (a specimen he named
Aublysodon molnaris, but which is now recognized as a juvenile Tyrannosaurus
rex), calling it Aublysodon huoyanshanensis and placing it Aublysodontinae
within the Tyrannosauridae. Though Paul's generic synonymy was not often followed,
his placement of Shanshanosaurus in an Aublysodontidae/inae was standard
through the 1990's, sometimes renamed Shanshanosaurinae (Olshevsky, 1995) due
to Aublysodon's indeterminate nature. Holtz (2001) was the first to include
Shanshanosaurus in a cladistic analysis, where it emerged as a basal
tyrannosaurine due to its low tooth count. However, Dong's tooth counts are
incomplete (Currie and Dong, 2001). Currie and Dong (2001) restudied and redescribed
the material, resulting in some corrections. The supposed postorbital identified
by Dong was a proximal rib, while the cervical vertebrae are amphicoelous, not
procoelous (contra Molnar et al., 1990). Indeed, nothing prevents the specimen
from being a juvenile tyrannosaurid, though Currie and Dong were reluctant to
assign it to any particular genus. They did note it was more similar to Tarbosaurus
than Alioramus in the arrangement of its maxillary nutrient foramina,
but Currie (2003) later indicated this was not diagnostic of Alioramus.
Carr (2005) found Shanshanosaurus emerged as the sister taxon to Tarbosaurus
+ Tyrannosaurus before ontogenetically influenced characters were taken
into account. Furthermore, he identified a synapomorphy present in Shanshanosaurus
and some Tarbosaurus individuals- a subcutaneous flange extending dorsally
off the horizontal maxillary ramus. Interestingly, some Tarbosaurus specimens
lack it (GIN coll., PIN 551-1, 553-1) and it's not ontogenetic. Perhaps sexual
or individual variation?
Non-Nemegt Tarbosaurus?- Although often touted as ranging widely
over Asia, diagnostic Tarbosaurus remains have only been verified from
the Nemegt Formation of Mongolia and (thanks to Shanshanosaurus) the
Subashi Formation of China. In addition to Shanshanosaurus, Dong (1977)
described some fragments from the latter locality found in 1964-1966 as Tarbosaurus sp..
Zhai et al. (1978) later listed the nomen nudum Tyrannosaurus "turpanensis",
which judging by the horizon, locality and known elements, is based on Dong's
material. These are provisionally referred to T. bataar here given its
presence in the formation and seeming absence of other tyrannosaurids in the
Nemegt (assuming the faunas are similar).
Fragmentary remains from the Quiba Formation of China were named Tyrannosaurus
luanchuanensis (Dong, 1979), later referred to Tarbosaurus (Olshevsky,
1991) and Jenghizkhan (Olshevsky, 1995). Carr and Williamson (2000) noted
its teeth have a denticle density like that of Tyrannosaurus, different
from Daspletosaurus and albertosaurines. Since Tarbosaurus has
the same density as Tyrannosaurus (Hurum and Sabath, 2003), and the Quiba
Formation may be contemporaneous with the Nemegt Formation, it is provisionally
considered a junior synonym of T. bataar.
Jerzykiewicz et al. (1993) referred premaxillary and maxillary teeth from the
Late Campanian Djadochta Formation of Mongolia to Tarbosaurus sp.. This
is slightly earlier than the Nemegt and Subashi Formations, suggesting they
are not from T. bataar at least.
Dong (1979) briefly described some unassociated elements from the Yuanpu (=Nanxiong)
Formation of China as Tarbosaurus sp.. As with the Djadochta material,
they are from Campanian deposits, suggesting they are not T. bataar even
if they are Tarbosaurus.
Nessov (1995) referred a femur (N 601/12457) from the Bostobe Formation of Kazakhstan
to Tarbosaurus sp. (incorrectly translated by Olshevsky, DML 1996 as
an ilium), but Carr (2005) determined it lacks the synapomorphies of Tarbosaurus
+ Tyrannosaurus and of Alectrosaurus. It seems to be a Beipiaosaurus-grade
therizinosaur. Bazhanov and Kostenko (1958) referred a dentary (IZK 33/MP-61)
from Kara-Cheku in Kazakhstan to Tyrannosaurus aff. bataar, but Averianov
et al. (2012) redescribed it as indeterminate within the Tarbosaurus-Tyrannosaurus
clade. Finally, Nessov noted tyrannosaurid remains in Bolotsky and Moisyeyenko
(1988) from the Udurchukan Formation of the Tsagayan Group of Russia, which
he stated were probably Tarbosaurus sp.. Bolotsky (2013) has referred
the larger teeth and metacarpal I from this formation to Tyrannosaurinae, but
characters of Tarbosaurus itself have yet to be identified.
Another specimen often referred to T. bataar (e.g. Molnar et al., 1990)
is Albertosaurus? periculosus from the Yuliangze Formation of China (equivalent
to the Udurchukan Formation). Bolotsky's thesis argued this and similar specimens
from the Udurchukan Formation were more likely albertosaurines based on their
abundance compared to larger specimens, but it remains possible these are juvenile
tyrannosaurines.
Chingkankousaurus fragilis is also sometimes listed as a junior synonym
of Tarbosaurus bataar, but is based on an indeterminate partial scapula
equally similar to other derived tyrannosauroids (Brusatte et al., 2013). It
is from the Wangshi Series of China, along with Tarbosaurus? zhuchengensis,
originally Tyrannosaurus zhuchengensis (Hu et al., 2001). Based on a
metatarsal and some referred teeth, the Chinese description has yet to be translated,
though the earlier age suggests it is not T. bataar.
Finally, Alioramus remotus from the Beds of Nogoon Tsav in Mongolia may
be a juvenile T. bataar, though this is is controversial (see entry).
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Tsogtbaatar, 2004. Fossil specimens prepared in Mongolian Paleontological Center 1993-2001. Hayashibara Museum of Natural
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Tsuihiji, Watabe, Witmer, Tsubamoto and Tsogtbaatar, 2007. A juvenile skeleton
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Brusatte, Carr, Erickson, Bever and Norell, 2009. A long-snouted, multihorned
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Tsuihiji, 2010. Reconstructions of the axial muscle insertions in the
occipital region of dinosaurs: Evaluations of past hypotheses on
Marginocephalia and Tyrannosauridae using the extant phylogenetic
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and Witmer, 2011. Cranial osteology of a juvenile specimen of Tarbosaurus
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T? sp. indet. (Gilmore, 1933)
Late Campanian, Late Cretaceous
Djadochta Formation, Inner Mongolia, China
Material- ?(AMNH 6522) (~8 m) partial ilium (Gilmore 1933)
premaxillary teeth, maxillary teeth (Jerzykiewicz et al., 1993)
Comments- The teeth were referred to Tarbosaurus sp.. by Jerzykiewicz,
which is possible though they are too early to be from T. bataar. Gilmore
(1933) described the ilium as a large theropod, perhaps a 'deinodontid'. It
is identified as a tyrannosaurid on the AMNH website. They are more likely to
be the contemporaneous Zhuchengotyrannus.
References- Gilmore, 1933. Two new dinosaurian reptiles from Mongolia
with notes on some fragmentary specimens. American Museum Novitates. 679, 1-20.
T? sp. indet. (Dong, 1979)
Campanian, Late Cretaceous
Yuanpu Formation of the Nanxiong Group, Guangdong, China
Material- (IVPP coll.; unassociated) third premaxillary tooth, lateral
tooth (72 mm), dorsal vertebra, fragmentary pedal elements (Dong, 1979)
Campanian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Material- (NHMG 8501) lateral tooth (~76 x 40.4 x 29 mm) (Mo and Xu,
2015)
Comments- Dong referred the IVPP remains to Tarbosaurus sp.. As
these are from earlier deposits than the Maastrichtian Nemegt and Subashi Formations,
they are probably not from Tarbosaurus bataar, and may not even belong
to this genus. Their serration density is similar to Tarbosaurus and
Tyrannosaurus, but not unequivocally different from large Campanian North
American teeth. They are more likely to be the contemporaneous Zhuchengotyrannus.
Mo and Xu (2015) described a tooth from Jiangxi which they referred to Tyrannosauridae
indet., noting it was comparable in size to Zhuchengotyrannus but more
recurved.
References- Dong, 1979. Cretaceous dinosaurs of Hunan, China. Mesozoic
and Cenozoic Red Beds of South China. In Institute of Vertebrate Paleontology
and Paleoanthropology and Nanjing Institute of Paleontology (eds.). Selected
Papers from the "Cretaceous-Tertiary Workshop". Science Press. 342-350.
Mo and Xu, 2015. Large theropod teeth from the Upper Cretaceous of Jiangxi,
southern China. Vertebrata PalAsiatica. 53(1), 63-72.
undescribed tyrannosaurine (Stein and Triebold, 2005)
Late Campanian, Late Cretaceous
Upper Judith River Formation, Montana, US
Material- (AMNH 30564) gastralium
....(RMDRC 02-001; Sir William; BCT) (~9.5 m; 1.76 tons; 15 year old subadult) lacrimal,
partial jugal, postorbital, squamosal, quadratojugal, ectopterygoid, pterygoid,
dentaries, cervical vertebrae, cervical ribs, dorsal vertebrae, dorsal ribs,
gastralia, fragmentary scapulocoracoid, ischia, femur (980 mm), fragmentary
tibia, fragmentary fibula, fragmentary astragalus
?(referred to lancensis) fifty teeth (Kemmick, 2004)
Comments- Discovered in 2001, this specimen was originally identified
as a young Tyrannosaurus rex and nicknamed Sir William (Anonymous, 2004).
It is listed as an individual of this species in Erickson et al. (2004) and
on the AMNH online catalogue. However, it later became clear it was preserved
in the Upper Judith River Formation, not the Hell Creek Formation (Stein and
Triebold, 2005). The latter authors believe this specimen represents a new taxon,
close to the ancestry of T. rex. The AMNH 30564 portion apparently consists
of a gastralium fragment, while the RMDRC reported the main specimen was being
prepared in their lab through March 2010 (Maltese, 2010 online).
Kemmick (2004) reported fifty Nanotyrannus teeth associated with this
specimen. Maltese (pers. comm., 2008) found these teeth were similar to albertosaurines
and Daspletosaurus in morphology.
References- Erickson, Makovicky, Currie, Norell, Yerby and Brochu, 2004.
Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs.
Nature, v. 430, p. 772-775.
Kemmick, 2004. T-rex roamed near Roundup: Fossil hunters stumbled across bones
2 years ago. The Billings Gazette. 9-4-2004, (pp ?].
Stein and Triebold, 2005. Preliminary analysis of a sub-adult tyrannosaurid
skeleton, known as "Sir William" from the Judith River Formation of
Petroleum County, Montana. The origin, systematics, and paleobiology of Tyrannosauridae.
27-28.
Maltese, 2010 online. BCT is finished! RMDRC paleo lab. 3-10-2020.
Tyrannosaurini Olshevsky, 1995
Tyrannosaurinae sensu Holtz, 2001
Definition- (Tyrannosaurus rex <- Aublysodon mirandus) (modified)
References- Olshevsky, 1995. The origin and evolution of the tyrannosaurids.
Kyoryugaku Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. In Tanke and
Carpenter (eds.). Mesozoic Vertebrate Life. 64-83.
Tyrannosaurus Osborn, 1905
= Manospondylus Cope, 1892 (nomen oblitum)
= Dynamosaurus Osborn, 1905
?= “Clevelanotyrannus” Bakker, Williams and Currie vide Currie, 1987
?= “Nanotyrannes” Carpenter vide Anonymous, 1988
?= Nanotyrannus Bakker, Williams and Currie, 1988
= Stygivenator Olshevsky, 1995
= Dinotyrannus Olshevsky, 1995
T. rex Osborn, 1905
= Manospondylus gigas Cope, 1892 (nomen oblitum)
?= Aublysodon amplus Marsh, 1892
?=Aublysodon cristatus Marsh, 1892
?= Deinodon amplus (Marsh, 1892) Hay, 1902
?= Deinodon cristatus (Marsh, 1892) Hay, 1902
= Dynamosaurus imperiosus Osborn, 1905
?= Tyrannosaurus amplus (Marsh, 1892) Hay, 1930
?= Gorgosaurus lancensis Gilmore, 1946
?= Deinodon lancensis (Gilmore, 1946) Kuhn, 1965
?= Aublysodon lancensis (Gilmore, 1946) Charig in Appleby, Charig, Cox,
Kermack and Tarlo, 1967
?= Albertosaurus lancensis (Gilmore, 1946) Russell, 1970
= Tyrannosaurus imperiosus (Osborn, 1905) Swinton, 1970
= Tyrannosaurus "vannus" Lawson, 1972
?= Manospondylus amplus (Marsh, 1892) Olshevsky, 1978
?= Nanotyrannus lancensis (Gilmore, 1946) Bakker, Williams and Currie,
1988
= Albertosaurus “megagracilis” Paul, 1988
= Aublysodon molnaris Paul, 1988
= Aublysodon molnari Paul, 1988 emend. Paul, 1990
= Tyrannosaurus “gigantus” Harlan, 1990
= Dinotyrannus megagracilis Olshevsky, 1995
?= Stygivenator amplus (Marsh, 1892) Olshevsky, 1995
?= Stygivenator cristatus (Marsh, 1892) Olshevsky, 1995
= Stygivenator molnari (Paul, 1988) Olshevsky, 1995
= Tyrannosaurus “stanwinstonorum” Pickering, 1995
= Tyrannosaurus “imperator” Melbourne, 1998
= Tyrannosauridae sensu Sereno, 1998
Definition- (Tyrannosaurus rex <- Alectrosaurus olseni, Aublysodon
mirandus, Nanotyrannus lancensis) (modified)
= Tyrannosaurus imperator Paul, Persons IV and Raalte, 2022
= Tyrannosaurus regina Paul, Persons IV and Raalte, 2022
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, North Dakota, South Dakota, Wyoming, US
Holotype- (CMN 9380; =AMNH 973) (12.4 m, 4.7 tons; adult) maxilla (695
mm), lacrimals, squamosal, ectopterygoid, dentaries (860 mm), surangular (610
mm), teeth, ninth cervical vertebra, second dorsal vertebra, eighth dorsal vertebra
(130 mm), ninth dorsal vertebra (145 mm), tenth dorsal vertebra, eleventh dorsal
vertebra, twelfth dorsal vertebra, thirteenth dorsal vertebra (170 mm), dorsal
ribs, three gastralia, sacrum (940 mm), scapula (950 mm), humerus (360 mm),
ilia (1.515 m), pubes (1.25 m), ischia (1.11 m), femur (1.28 m), tibia (1.14
m), metatarsal I, metatarsal II (615 mm), distal metatarsal III (~684 mm), metatarsal
IV (600 mm), phalanx IV-1
Referred- (AMNH 1011) incomplete tooth (Molnar, 1991)
(AMNH 5005) (juvenile) cranial fragments, femur (not collected), fibula (Molnar,
1991)
(AMNH 5020) metatarsal IV (Molnar, 1991)
(AMNH 5021) pedal phalanx (Molnar, 1991)
(AMNH 5027) (12.4 m, 5.7 tons, adult) skull (1.355 m; maxillae 710 mm), mandibles
(1.205 m; dentary 850 mm), cervical vertebrae 1-10 (960 mm total), nine cervical
ribs, (dorsal series 2.184 m) dorsal vertebrae (~160 mm), twelfth dorsal vertebra
(161 mm), thirteenth dorsal vertebra, twenty dorsal ribs, sacrum, first caudal
vertebra, caudal vertebrae 1-15, 17, 21, 22, seven chevrons, ilia (1.515 m),
pubes (~1.2 m), ischia (1.236 m) (Osborn, 1912)
(AMNH 5044) caudal vertebrae (Molnar, 1991)
(AMNH 5050) partial dentary (Osborn, 1916)
(BHI 3033; Stan) (12.3 m; 3.7 tons; adult) skull (~1.4 m; maxilla 775 mm), mandibles
(1.34 m; dentary 915 mm), thirty-five teeth, ten cervical vertebrae, fourteen
cervical ribs, thirteen dorsal vertebrae, twelve dorsal ribs, sacrum (1.06 m),
thirty-one caudal vertebrae, twenty-four chevrons, ilia (1.55 m), proximal pubes,
proximal ischia, femora (1.31 m), tibiae, fibula, astragali, calcanea, metatarsal
II (595 mm), metatarsal III, metatarsal IV (600 mm), eleven pedal phalanges
(Larson, 1992)
(BHI 4100; Duffy) (subadult) incomplete skull (premaxilla, maxillae (730 mm),
nasals, lacrimals, jugals, postorbital, squamosal, quadratojugals, quadrates,
palatines, ectopterygoid, pterygoid, epipterygoid, partial braincase), incomplete
mandible (dentary (770 mm), splenial, coronoid, surangular, prearticular), dentary,
forty-nine teeth, thirteen presacral vertebrae, nine dorsal ribs, eight caudal
vertebrae, six chevrons, scapulae (800 mm), coracoids, ischium, astragalus (Browne,
1993)
(BHI 4182; Fox or County rex) postorbital, quadratojugal, ectopterygoid, mandibles
(dentary 910 mm), forty-three teeth, two cervical vertebrae, two cervical ribs,
dorsal vertebra, five dorsal ribs, three caudal vertebrae (Larson, 2008a)
(BHI 6219; 007) premaxillae, maxillae, partial dentary, vertebra, dorsal rib,
distal humerus, partial tibia, partial fibula, metatarsal, pedal phalanx (Larson,
2008a)
(BHI 6249; Steven) two incomplete cranial elements, six incomplete dorsal vertebrae,
five dorsal ribs, incomplete femur, phalanx, eggshells (Larson and Donnan, 2002)
?(BHI 6235; referred to lancensis) (juvenile) lacrimal?, jugal, frontal,
three teeth (Erickson, 1999)
(BHI coll.) (subadult) proximal tibia, fibula (Erickson, 1999)
(BHI coll.; Rex B; Triceratops Alley rex) premaxilla, maxilla, nasals, lacrimals,
frontals, quadratojugal, quadrate, braincase, ectopterygoid, rib, scapula, coracoid
(Larson, 2008a)
?(BMRP 2002.4.1; Jane; referred to lancensis) (~6.4 m; ~680 kg; 11 year
old juvenile) incomplete skull (724 mm; maxilla 470 mm), mandible (dentary 505
mm), teeth (~100 mm), seven cervical vertebrae, cervical ribs, four posterior
dorsal vertebrae, dorsal ribs, gastralia, sacrum (500 mm), twenty proximal caudal
vertebrae (~130 mm), seventeen chevrons, scapulocoracoid, humerus (280 mm),
radius, ulna, partial manus, ilia (720 mm), pubes, ischia, femora (720 mm),
tibiae (840 mm), metatarsal II (510 mm), metatarsal IV (513 mm), phalanx III-1
(135 mm), phalanx III-2 (103 mm), pedal phalanges (Henderson, 2005)
?(BMRP 2006.4.4; Petey) (~7-7.4 m; 10 year old juvenile) five or six dorsal
and caudal vertebrae, more than four dorsal ribs, gastralia, scapulocoracoid,
humerus, partial (?)ulna, (?)metacarpal fragments, two manual ungual I, manual
ungual II, femur, partial tibia, fibula, astragalus, pedal ungual I, eight pedal
phalanges (Tremaine et al., 2014)
(CMI 2001.90.1; = BHI 4960; Bucky) (10 m; 2.98 tons; 16 year old adult) cervical
vertebrae 3-10, eleven cervical ribs, nine dorsal vertebrae, sixteen dorsal
ribs, twenty-four gastralia, sacrum (895 mm), five proximal caudal vertebrae,
three mid caudal vertebrae, six distal caudal vertebrae, fourteen chevrons,
scapulae (940 mm), coracoid, furcula, ulna (176 mm), manual phalanx I-1, metacarpal
II, ilia (1.275 m), ischium, (femur ~1.168 m) metatarsal II (550 mm), pedal
phalanx II-1, pedal phalanx II-2, pedal ungual II, pedal phalanx III-3, metatarsal
IV (565 mm), pedal phalanx IV-2, pedal phalanx IV-4, metatarsal V (Larson and
Rigby, 2005)
?(CMN 7541; holotype of Gorgosaurus lancensis) (juvenile) skull (602
mm; maxilla 385 mm), mandibles (dentary 375 mm) (Gilmore, 1946)
(CMN coll.; Mr. Zed; = Z-rex; = Samson) (~12.6 m) skull (1.4 m), mandibles (dentary
870 mm), twenty-two teeth, nine cervical vertebrae, two cervical ribs, seven
dorsal vertebrae, ten dorsal ribs, seventeen caudal vertebrae, four chevrons,
femora (1.295 m), tibial fragments, fibula, metatarsal II (610 mm), metatarsal
III, metatarsal IV (635 mm), ten pedal phalanges (Glut, 2002)
?(DDM 35) (juvenile) frontal, partial ?radius, tibia, pedal phalanx (Carr, de
Santis, Wojahn, Brown and Ogle, 2007)
(FMNH PR2081; =BHI 2033; Sue; material of Tyrannosaurus "gigantus";
material of Tyrannosaurus "stanwinstonorum"; holotype of Tyrannosaurus imperator) (12.8 m; 5.654
tons; 28 year old adult) skull (1.394 m; maxilla 861 mm), stapes, mandibles
(1.437, 1.395 m; dentary 1.01 m), proatlas arches, axis (142 mm), third cervical
vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth cervical
vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical
vertebra, axial ribs, twelve cervical ribs (350-610 mm), 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, nineteen dorsal
ribs (.737-1.473 m), gastralia, sacrum (948 mm), thirty-six caudal vertebrae,
twenty-five chevrons, scapulocoracoids (1.303, 1.310 m; scapula 1.14 m), furcula,
humerus (385 mm), radius (173 mm), ulna (214 mm), metacarpal I (65 mm), phalanx
I-1 (75 mm), manual ungual I, metacarpal II (104 mm), phalanx II-1 (45 mm),
manual ungual II, ilia (1.46 m), pubes, ischia, femora (1.321, 1.308 m), tibiae
(1.143 m), fibulae (1.029, 1.035 m), astragali, calcanea, distal tarsal IV,
pedal ungual I, metatarsal II (584 mm), phalanx II-1 (207 mm), phalanx II-2
(152 mm), pedal ungual II (175 mm), metatarsal III (671 mm), phalanx III-1 (201
mm), phalanx III-2 (136 mm), phalanx III-3 (122 mm), pedal ungual III (204 mm),
metatarsal IV (621 mm), phalanx IV-1 (154 mm), phalanx IV-2 (111 mm), phalanx
IV-3 (88 mm), metatarsal V (275 mm) (Brochu, 2003)
(FMNH PR2902) premaxillary tooth (12.2x4.8x2.9 mm) (Gates et al., 2015)
(Great Plains Paleontology coll.; Rex A; Ollie) premaxillae, maxilla, jugal,
postorbitals, quadrates, partial braincase, pterygoids, several cervical vertebrae,
cervical ribs, dorsal vertebrae, dorsal ribs, caudal vertebrae, several chevrons,
scapula, humeri, radius, ulna, ilium, pubis, ischium, femora, tibiae, fibulae,
astragali, calcanea, two metatarsals, several phalanges (Larson, 2008a)
(Great Plains Paleontology coll.; Otto) cervical ribs, dorsal ribs, caudal vertebrae,
femora, tibiae, fibula, two metatarsals (Larson, 2008a)
(HMNS 2006.1743.01; = BHI 6230; Wyrex) (11.8 m; 3.6 tons) maxilla, jugal, partial
postorbital, partial squamosal, quadratojugal, partial quadrate(?), partial
pterygoid, basioccipital, exoccipital-opisthotic, partial surangular, angular,
partial prearticular, articular, atlas, cervical vertebra, five cervical ribs,
five dorsal vertebrae, fifteen partial dorsal ribs, seventeen gastralia, incomplete
sacrum, eleven caudal vertebrae, more than four chevrons, scapula, coracoid,
humerus (330 mm), ulna (185 mm), radiale, metacarpal I, metacarpal II, metacarpal
III, ilium (1.47 m), pubis, ischia, femora (1.19 m), tibia, fibulae, astragalus,
calcaneum, distal tarsal III, distal tarsal IV, phalanx I-1, metatarsal II (600
mm), phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1,
phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (625 mm), phalanx
IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, metatarsal V, skin impressions
(Larson, 2008a)
(LACM 23844) (adult) incomplete skull, mandibles (1.39 m- dentary 920 mm), two
cervical vertebrae, seven dorsal vertebrae, five dorsal ribs, gasteralia, four
caudal vertebrae, ten chevrons, scapula, incomplete ischia, femur, tibia, astragalus,
metatarsus (640 mm), ten pedal phalanges (Molnar, 1991)
(LACM 23845; holotype of Albertosaurus megagracilis) (~9.6 m, ~1.81 tons,
14 year old subadult) partial skull (900 mm), partial mandibles, scapula, coracoid,
ulna (131 mm), metacarpal II (70 mm), proximal femur (~989 mm), proximal tibia,
fibula (863 mm), astragalus, pedal ungual I, metatarsal II (507 mm), phalanx
II-1, phalanx II-2, distal metatarsal III, phalanx III-1, phalanx III-2, phalanx
III-3, pedal ungual III, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4,
pedal ungual IV (Molnar, 1980)
(LACM 28471; Jordan theropod; holotype of Aublysodon molnari) (~2.5 m;
30 kg; 2 year old juvenile) (skull ~450 mm) premaxillary tooth (12 mm), partial
maxillae, maxillary teeth, nasals, frontals, parietals, partial dentary, dentary
teeth, six teeth (24-32 mm), surangular fragment (femur ~252 mm) (Molnar, 1978)
(LACM 7509/150167; Thomas) maxillae, lacrimal, jugals, frontals, postorbital,
squamosal, quadratojugal, quadrate, braincase, ectopterygoid, dentaries, posterior
mandibular elements, 30-35 teeth, few dorsal vertebrae, ribs, gastralia, sacrum,
about twenty caudal vertebrae, scapulae, coracoids, ilia, ischia, femora, tibiae,
fibulae, astragali, calcanea, metatarsi, pedal phalanges, unprepared elements
(Larson, 2008a)
(MOR 008) (~13.8 m?) incomplete skull (missing premaxilla, vomer, palatine and
epiterygoid) (1.50 m; maxilla 720 mm), incomplete mandibles (dentary 880 mm),
atlas (Molnar, 1991)
(MOR 009; = GE-69-1; Hager rex) (11.1 m) maxilla, partial jugal, partial lacrimal,
frontal, postorbital, partial squamosal, dentary, teeth, dorsal vertebrae, four
dorsal ribs, twenty-two caudal vertebrae, seven chevrons, ilia (1.16 m), pubes,
ischia, femora (1.143 m), incomplete tibiae (1.118 m), fibula, astragalus, metatarsus
(593 mm), seven pedal phalanges (Larson, 2008a)
(MOR 557) posterior braincase (MOR online)
(MOR 980; Rigby specimen; Peck's rex; material of Tyrannosaurus "imperator")
(~12.8 m; 3.4 tons; adult) incomplete skull (~1.37 m; maxilla 770 mm), partial
mandibles (dentary 900 mm), cervical vertebrae, cervical ribs, dorsal vertebrae,
several dorsal ribs, gastralia, sacrum (851 mm), nine or ten proximal caudal
vertebrae, proximal chevrons, scapulae (940 mm), coracoid, furcula, humeri (362
mm), metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-?,
manual ungual II, metacarpal III (~254 mm), ilia (1.397 m), pubes (~1.32 m),
ischia, femur (1.232 m), tibia, fibula, astragalus, calcaneum, metatarsal II
(597 mm), metatarsal IV (655 mm) (Larson and Rigby, 2005)
(MOR 1127; L-rex) cervical vertebrae, cervical ribs (MOR online)
(MOR 2925) 29 postcranial elements including eight vertebrae including atlas,
seven ribs and pubis (Hall and Keenan, 2010)
(MOWT L07-023-001; Ivan) about fifteen presacral vertebrae, about twenty-five
presacral ribs, gastralia, sacrum, about twenty-five caudal vertebrae, about
thirty chevrons, scapulocoracoid, partial ilia, pubes, ischia, femur, tibia,
fibula, astragalus, two metatarsals, six pedal phalanges (Larson, 2008a)
(North Carolina Museum of Natural Sciences coll.; Dueling Dinosaurs in part; referred to lancensis) (juvenile)
incomplete skull, mandibles, hyoid, cervical vertebrae, cervical ribs, dorsal
vertebrae, dorsal ribs, gastralia, partial scapula, humeri (~268 mm), radii, ulnae (~204 mm), radiale,
distal carpal I, distal carpal II, metacarpals I (~49 mm), phalanges I-1, manual unguals
I, metacarpal II (~92 mm), phalanx II-1 (~51 mm), phalanges II-2, manual unguals II, metacarpal
III, phalanx III-1, ilium, pubes, proximal ischium, femur, partial tibia, metatarsal
I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal
ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal
ungual III, partial metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal ungual IV, skin impressions (http://www.bonhams.com/auctions/21076/lot/1032/)
(NHMUK R7995; = AMNH 5881) gastralia, femur, tibiae, fibula?, metatarsal I, metatarsal
II, metatarsal IV, pedal phalanges? (Osborn, 1906)
(RGM 792.000) (robust adult) skull, incomplete axial column including proximal
and mid caudal vertebrae, dorsal ribs, chevrons, scapula, coracoid, furcula,
pelvis and hindlimb including femur (Schulp, Bastiaans, Kaskes, Manning and
Larson, 2015)
?(RMDRC coll.; referred to lancensis) premaxillary tooth, four lateral teeth (Maltese, 2013 online)
(SDSM 8354/PRB8775) skull, partial skeleton (Carpenter pers. comm. to Ford and
Chure, 2001)
(SDSM 12047; Mud Butte T. rex) (subadult) skull lacking premaxilla, dentaries,
coronoid, angular, three partial ribs, caudal vertebrae 15-34, chevrons (Bjork,
1985)
(SDSM 64351) tooth (Stokosa, 2005)
(Trails Regional Museum coll.; Bowman) 45 elements including vertebrae, ribs,
gastralia, distal scapula and pubes (Oakland and Pearson, 1995)
(UCMP 118742) (~12.1-12.4 m, adult) (skull ~1.31 m?) maxilla (810 mm) (Molnar,
1991)
(UCMP 124488) tooth (UCMP online)
(UCMP 131583) maxilla, dentaries, postcranial fragments (Molnar, 1991)
(UCMP 136518) partial femur (Hutchinson, 2001)
(UCMP 137537) incomplete pes (UCMP online)
(UCMP 137538) phalanx (UCMP online)
(UCMP 137539) incomplete pes (Snively and Russell, 2003)
(UCMP 137540) incomplete pes (UCMP online)
(UCMP 137541) metatarsal (UCMP online)
(UCMP 137542) phalanx (UCMP online)
(UCMP 140418) humerus (UCMP online)
(UCMP 140506) vertebra, ribs, ilium, ischium fragments (UCMP online)
(UCMP 140600) tooth (UCMP online)
(UCMP 154426) tooth (UCMP online)
(UCMP 154585) distal metatarsal (UCMP online)
(UCMP 154586) metatarsal fragments (UCMP online)
(UCMP 172032) tooth (UCMP online)
(UCMP 172228) tooth fragments (Holroyd and Hutchison, 2002)
(UCMP 172366) tooth fragment (Holroyd and Hutchison, 2002)
(UND-PC 15840) fragmentary tooth (Hoganson and Murphy, 2002)
(University of Illinois coll.) tooth (Jacobson and Sroka, 1995)
(USNM 555000; = MOR 555; Wankel rex; holotype of Tyrannosaurus regina) (12.4 m; 4.0 tons; adult) incomplete skull (maxilla 798
mm), dentary (990 mm), cervical vertebrae 2-10, dorsal vertebrae 1-13, sacrum
(1.01 m), caudal vertebrae 1-18, scapulae (980 mm), coracoids, humerus (377
mm), radius, ulna (198 mm), semilunate carpal, metacarpal I, phalanx I-1 (98
mm), metacarpal II (94 mm), phalanx II-1 (57 mm), phalanx II-2 (78 mm), metacarpal
III, ilia (1.49 m), pubes, ischia, femora (1.275 m), tibiae (1.1 m), metatarsal
II (585 mm), metatarsal II (657 mm), metatarsal IV (605 mm), pedal phalanges
(Horner and Lessem, 1993)
(USNM coll.; Nathan or N-rex) incomplete dentary, angular, cervical vertebra,
two cervical ribs, two dorsal neural spines, two dorsal ribs, gastralium, three
caudal vertebrae, three chevrons, ilium, pubis, ischium, femur, tibia, fibula,
pes (Larson, 2008a)
(UWGM 181) maxilla, jugal, postorbitals, squamosal, quadratojugal, quadrate,
partial braincase, partial pterygoid, dentaries, splenial, surangular, prearticular,
three vertebrae, 100 fragments (Larson, 2008a)
(YPM 8228) (YPM online)
(private coll.; Tinker) (~8 m; subadult) premaxillae, maxillae, partial nasal,
jugal, parietal, squamosal, quadratojugals, quadrate, palatine, pterygoid, dentary,
splenial, coronoids, surangular, angular, preartcular, articulars, teeth, two
cervical ribs, five dorsal ribs, rib fragments, twenty partial caudal vertebrae,
twelve chevrons, partial scapulae, coracoid, humeri, manual ungual, incomplete
ilia, pubes, ischium (650 mm), tibia (670 mm), pedal ungual (Larson, 2008a)
(private coll.; referred to lancensis) (juvenile) teeth (with Tinker)
(Blasing, DML 2006)
(private coll.; Belle) (subadult) (Blasing, DML 2006)
(private coll.; Regina) (adult) (with Tinker) maxilla, jugal (Blasing, DML 2006)
(private coll.; Rex C) premaxilla, maxilla, splenial, surangular, articular,
cervical vertebra, dorsal vertebra, two caudal vertebrae, chevron, ischium,
tibia, fibulae, astragalus, three pedal phalanges, fragments (Larson, 2008a)
(private coll.; Wayne) dorsal vertebra, several rib or gastralia fragments,
nineteen caudal vertebrae, two chevrons, elements (Larson, 2008a)
five teeth, vertebral fragment, distal metatarsal, phalanx (Lupton, Gabriel
and West, 1980)
?(referred to lancensis) (juvenile) phalanx (Stenerson and O'Conner,
1994)
(partly referred to lancensis) cranial elements, teeth, distal caudal
vertebrae, chevrons, phalanges (DePalma, 2010)
(juvenile) dorsal vertebrae, ribs, gastralia, scapulocoracoid, humerus, ulna,
manual unguals, femur, tibia, fibula, pedal ungual (Williams, Brusatte, Mathews
and Currie, 2010)
(juvenile) caudal vertebra, humerus, pedal elements (Holtz, Williams, Tremaine
and Matthews, 2014)
Late Maastrichtian, Late Cretaceous
Denver Formation, Colorado, US
(DMNH 2827) (10.8 m) three teeth, ribs, distal caudal vertebra, scapula (820
mm), coracoid (240 mm), partial ilium (~1.85 m), incomplete femur (~1.11 m),
distal tibia, fibula (872 mm), astragalus (288 mm wide) (Carpenter and Young,
2002)
(DMNH 32825) tooth (Carpenter and Young, 2002)
(UCMP 36303) tooth (Carpenter and Young, 2002)
(UCMP 38804) tooth (Carpenter and Young, 2002)
(YPM 4192) tooth (Carpenter and Young, 2002)
? mandible (Cannon, 1888)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
material (Wroblewski, 1998)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
(RSM 2523.8; Scotty) incomplete skull, incomplete mandibles, more than forty
cervical, dorsal and caudal vertebrae, sixteen dorsal ribs, scapula, manual
phalanx, ilia, pubes, ischia, femur (1.29 m), tibia, fibula, metatarsal, several
pedal phalanges (Tokaryk and Bryant, 2004)
pedal phalanges (Langston’s 1955 field notes in Ford and Chure 2001)
Early Maastrichtian, Late Cretaceous
Lower Hell Creek Formation, Montana, South Dakota, US
(BHI 6248; E. D. Cope) maxilla, ectopterygoid, dentary, angular, cranial elements,
vertebrae, ribs (Larson, 2008a)
(MOR 1125; B-rex; Bob) (~10.4 m; 3.9 tons; 18 year old adult female) incomplete
skull missing premaxillae (maxilla 680 mm), mandibles missing a dentary (dentary
760 mm), three cervical vertebrae, four cervical ribs, four dorsal vertebrae,
thirteen dorsal ribs, sacrum, twelve caudal vertebrae, seven chevrons, scapulocoracoid,
furcula, ulna (200 mm), femora (1.07 m), tibiae, fibulae, astragalus, calcaneum,
eleven pedal phalanges (Schweitzer et al., 2005a)
(MOR 1126; Celeste or C-rex) (~14.1 m?) surangular, prearticular, three partial
dorsal vertebrae, twenty dorsal ribs, chevron (Larson, 2008a)
(MOR 1128; G-rex) (5.6 tons) incomplete dentary, two teeth, four dorsal vertebrae,
seven ribs, caudal vertebra, three chevrons, partial scapula, pubes, ischia,
femur (1.26 m), tibia (Larson, 2008a)
(MOR 1131; J-rex) cranial elements including frontals, parietals, braincase
(Larson, 2008a)
(MOR 1152; Frank or F-rex) posterior dorsal vertebrae, posterior dorsal ribs,
seven caudal vertebrae, four chevrons, pelvis, hindlimb, metatarsal (Larson,
2008a)
?(juvenile and adult) ninety-one teeth (Larson, Nellermoe and Gould, 2003)
Late Maastrichtian, Late Cretaceous
Javelina Formation, Texas, US
(BIBE 45850) (subadult) partial tibia (837 mm), partial pes (Wick, 2014)
(TMM 41436-1; material of Tyrannosaurus "vannus") (subadult)
maxilla (Lawson, 1976)
(TMM 46028-1) incomplete ~eleventh caudal vertebra (173 mm) (Wick, 2014)
Late Maastrichtian, Late Cretaceous
Lance Formation, Montana, South Dakota, Wyoming, US
(AMNH 3982; holotype of Manospondylus gigas) tenth cervical centrum (90
mm), cervical centrum (lost) (Cope, 1982)
(AMNH 5117) (adult) braincase, postorbital, pterygoid, hyoid (lost) (Osborn,
1912)
(BIOPSI coll.; Monty) premaxilla, maxilla, nasals, lacrimal, jugal, postorbital,
squamosal, quadratojugal, quadrates, braincase, pterygoids, surangular, four
cervical vertebrae, two dorsal vertebrae, twelve dorsal ribs, four gastralia,
three caudal vertebrae, (?)ulna, partial ilium, pubis, pedal phalanx, several
elements (Larson, 2008a)
(CMN 244) pedal phalanx (Molnar, 1991)
(CMN 1400) premaxilla, maxilla (760 mm), nasals, braincase, pterygoid, two cervical
ribs, dorsal vertebra, dorsal rib, three chevrons, pubic fragments, ischial
fragments (McIntosh, 1981)
....(CMN 9401) fragmentary lacrimal (Molnar, 1991)
(CMN 9379; =AMNH 5029) braincase, splenial (lost), prearticular (lost), articular
(lost) (Osborn, 1912)
(DIS 101) fragmentary skull, fragmentary skeleton (Anonymous, 1997)
(DMNH coll.) (juvenile) five teeth (Bakker et al., 1988)
?(DMNH coll.; referred to lancensis) (juvenile) three teeth (Bakker et
al., 1988)
(LDP 977-2; Pete) (9.4 m) anterior cervical vertebra, five posterior cervical
vertebrae, cervical rib, five anterior dorsal vertebrae (second dorsal vertebra
110 mm), presacral vertebrae, ten dorsal ribs, dorsal rib fragments, four gastralia,
gastralia fragments, two proximal caudal vertebrae, distal caudal vertebra,
scapular fragments, shaft of hindlimb element (Derstler and Myers, 2008a)
(MMS 51-2004) frontal, partial braincase (Molnar, 1978)
(NHMUK R7994; holotype of Dynamosaurus imperiosus; = AMNH 5866) (~11.5
m; 3.5 tons) palatines (lost), dentaries, atlas (65 mm), axis (100 mm), third
cervical vertebra (100 mm), fouth cervical vertebra (120 mm), fifth cervical
vertebra (115 mm), sixth cervical vertebra (120 mm), seventh cervical vertebra
(110 mm), eighth cervical vertebra (125 mm), ninth cervical vertebra (100 mm),
tenth cervical vertebra (110 mm), thirteen cervical ribs, first dorsal vertebra
(100 mm), second dorsal vertebra, third dorsal neural spine, fourth dorsal centrum,
fifth dorsal centrum, two sacral vertebrae, sacral neural spine, fragmentary
ilium, ischium, fragmentary femur (Osborn, 1905)
(SDSM 15115) posterior premaxillary tooth fragment (Whitmore, 1988)
(SDSM 15117) tooth fragment (Whitmore, 1988)
?(SDSM 15135) tooth tip (Stokosa, 2005)
?(SDSM 64287) posterior tooth (Stokosa, 2005)
(TATE coll.; Lee Rex) thirty-four elements including cervical ribs, dorsal ribs
and femur (Dalman, 2013)
(UCMP 73081) (UCMP online)
(UCRC PV1) (~8.5 m) presacral vertebrae, dorsal ribs, gastralia, scapulocoracoids,
coracoid fragments, furcula, forelimbs, hindlimb fragments (Lipkin and Sereno,
2004; described in Lipkin et al., 2007)
(USNM 2110) (~12.2 m) distal metatarsal IV (~590 mm) (Marsh, 1890)
(USNM 6183) (~9.8 m; 2.4 tons) femur (1.033 m), tibia (890 mm), proximal fibula
(Marsh, 1892)
(USNM 8064) ilium (Marsh, 1892)
?(YPM 296; holotype of Aublysodon amplus) (juvenile) premaxillary tooth
(27 mm) (Marsh, 1892)
?(YPM 297; holotype of Aublysodon cristatus) (juvenile) premaxillary
tooth (Marsh, 1892)
(YPM 1866) (YPM online)
(YPM 2220) premaxillary tooth (48x16.1x15.3 mm), maxillary tooth (40x20x10.1
mm) (Dalman, 2013)
(YPM 54459) tooth (16x10.2x8 mm), tooth (40x18x10.2 mm), tooth (?x22.2x12.1
mm), tooth (20x10x2.1 mm), tooth (22x10.2x2.1 mm), tooth (30x13x11 mm), tooth
(32.5x12.2x10.5 mm) (Dalman, 2013)
(YPM 57488) pedal phalanx II-2 (165 mm), phalanx III-2 (125 mm) (Dalman, 2013)
(YPM-PU 16516) anterior dentary (Dalman, 2013)
(YPM-PU 18307) (YPM online)
(YPM-PU 21203) (YPM online)
tooth fragments (Estes, 1964)
tooth (Browne, 1992)
partial tooth (Ein, 1993)
fragmentary teeth (Ein, 1993)
(commercial coll.) dorsal vertebrae (Derstler, 1994)
(private coll.) pedal elements (Derstler, 1994)
(juvenile) distal metatarsal (Derstler, 1994)
teeth (Derstler, 1995)
?(referred to lancensis) (juvenile) teeth (Derstler, 1995)
teeth (Spencer et al., 2001)
?(juvenile; referred to lancensis) teeth (Spencer et al., 2001)
(private coll.; Barnum) premaxillary fragment, two premaxillary teeth, maxillae,
maxillary tooth, jugal, squamosal, ectopterygoid, partial braincase, partial
dentary, three dentary teeth, surangular, angular, cervical vertebra, four dorsal
vertebrae, nine dorsal ribs, gastralia, three caudal vertebrae, partial scapula,
partial humerus, manual ungual, partial ilium, pubes, partial ischium, femora,
tibia, partial fibula, astragalus, calcaneum, partial metatarsal I, metatarsal
II, partial metatarsals III, phalanx III-1, metatarsal IV, phalanx IV-3, phalanx
IV-4 (Larson, 2008a)
Late Maastrichtian, Late Cretaceous
Livingston Formation, Montana, US
material (McMannis, 1965)
Maastrichtian, Late Cretaceous
Lomas Coloradas Formation of the Cabullona Group, Mexico
Material- (ERNO 8549) tooth (71 x 33 x 24 mm) (Serrano-Bra�as, Torres-Rodr�guez,
Reyes Luna, Gonz�lez and Gonz�lez-Le�n, 2014)
(ERNO 8550) tooth (41 x 33 x 19 mm) (Serrano-Bra�as, Torres-Rodr�guez,
Reyes Luna, Gonz�lez and Gonz�lez-Le�n, 2014)
(ERNO 8551) tooth (35 x 22 x 17 mm) (Serrano-Bra�as, Torres-Rodr�guez,
Reyes Luna, Gonz�lez and Gonz�lez-Le�n, 2014)
(ERNO 8552) tooth (39 x 27 x 19 mm) (Serrano-Bra�as, Torres-Rodr�guez,
Reyes Luna, Gonz�lez and Gonz�lez-Le�n, 2014)
(ERNO 005) (juvenile?) tooth (28 x 16 x 9 mm) (Serrano-Bra�as, Torres-Rodr�guez,
Reyes Luna, Gonz�lez and Gonz�lez-Le�n, 2014)
(ERNO 006) (juvenile?) tooth (37 x 21 x 12 mm) (Serrano-Bra�as, Torres-Rodr�guez,
Reyes Luna, Gonz�lez and Gonz�lez-Le�n, 2014)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
(NMC 9554) incomplete cervical vertebra (Russell, 1970)
(RTMP 81.12.1, including NMC 9950; Huxley rex) (12.5 m; 5.04 tons; 22 year old
adult) postorbital, seven anterior dorsal vertebrae, dorsal rib, partial sacrum
(980 mm), eight proximal caudal vertebrae, five proximal chevrons, ilia, pubis,
ischium, femora (1.284 m), tibiae (1.18 m), fibulae, astragalus, calcaneum,
distal tarsal III, distal tarsal IV, metatarsal III (698 mm), pedal phalanx
IV-1 (53 mm), six pedal phalanges (Russell, 1970)
(uncollected) skull (Currie pers. comm. to Ford and Chure 2001)
Late Maastrichtian, Late Cretaceous
Hall Lake Member of the McRae Formation, New Mexico, US
(NMMNH P-3698; = NMMNH P-1013-1) postorbital, squamosal, palatine (missidentified
as an articular), dentary, splenial, prearticular, articular, three teeth, three
chevrons (Gillette, Wolberg and Hunt, 1986)
Late Maastrichtian, Late Cretaceous
North Horn Formation, Utah, US
?(UMNH 7515) ungual (Difley and Ekdale, 2002)
(UMNH 7626) partial tooth (Difley and Ekdale, 2002)
(UMNH 11000) postorbital, squamosal, third cervical vertebra, fourth cervical
vertebra, dorsal rib, second sacral vertebra, third sacral vertebra, fourth
sacral vertebra, six mid caudal vertebrae, six chevrons, partial ilium, proximal
ischium, tibia, fibula, astragalus (Sampson and Loewen, 2005)
Late Maastrichtian, Late Cretaceous
Willow Creek Formation, Alberta, Canada
(RTMP 81.6.1; Black Beauty) (11.7 m; 3.23 tons; 18 year old adult) skull, partial
mandibles (dentary 770 mm), five cervical vertebrae, two cervical ribs, seven
dorsal vertebrae, eight dorsal ribs, humerus (302 mm), manual phalanx, femora
(1.21 m), tibiae, fibula, astragalus, calcaneum, four metatarsals, five pedal
phalanges (Currie, 1993)
Late Cretaceous
Alberta, Canada
?(referred to lancensis) skull, skeleton (Langston's 1955 field notes;
www.paleofile.com)
Late Cretaceous
Saskatchewan, Canada
pedal phalanx (Langston's 1955 field notes; www.paleofile.com)
Late Cretaceous
Montana, US
(MOR 1156; J-rex2) four elements (MOR online)
(MOR 1190) phalanx (MOR online)
(MOR 1191) fibula (MOR online)
(MOR 1198; Jen-rex) femoral fragment, phalanx (MOR online)
(MOR 1602; H-rex) pedal phalanx (MOR online)
(MOR 1628) maxilla (MOR online)
(private coll.; Cupcake) (subadult) skull, mandibles (Carr, online 2004)
(private coll.; King Kong) (~12 m) ~65% complete specimen including forelimbs
(Carr, online 2004)
(private coll.; Tristan) (~12 m) incomplete skull, postcrania including pectoral
girdle and pelvis (Stemmler, online 2015)
(private coll.; Russell) (two adults) partial skeletons (Express Newspapers, online 2013)
?
(AMNH 21542) (juvenile) partial dentary (Carr, 1999)
(BHI 116) frontal (Currie, 2003)
(BHI 1281) tooth (90 mm)
(BHI 6231) humerus (360 mm) (Larson, 2008)
(BHI 6232) (4.3 tons) femur (1.18 m) (Larson, 2008)
(BHI 6233) (4.1 tons) femur (1.11 m) (Larson, 2008)
(BHI 6242; Henry) (4.0 tons) femur (1.18 m) (Larson, 2008)
(LL 12823) (3.1 tons) femur (1.20 m) (Larson, 2008)
(RSM 283.2) frontal (Currie, 2003)
(RTMP 82.50.11) maxilla (Molnar, 1991)
(UCMP 154587) fibula (UCMP online)
Diagnosis- (after Carr, 2005) lacrimal horn absent; anterior margin of
dorsal quadratojugal process is notched; dorsolateral process of palatine inflated;
less than fifteen dentary teeth in adults.
Comments- Although often said to be known from few specimens in popular
works, a large number of fairly complete specimens are known, with more being
discovered each year and most remaining undescribed. This is no doubt due to
the extensive fieldwork done in the Hell Creek and Lance Formations, the distinctive
nature and size of Tyrannosaurus remains, and the popularity of the animal.
In general, specimens discovered since 1990 have not been described in the technical
literature. Because of its fame, many specimens get nicknames and are reported
to the popular press prior to mention in the technical literature, so that original
references are more difficult to track down. Thus the references given in the
materials list below may not be the first published. Osborn (1916) questionably
referred AMNH 5050 to Ornithomimus velox, but it is a tyrannosaurid dentary,
probably Tyrannosaurus itself based on provenance.
Lance Aublysodon species- Marsh (1892) described two new species
of Aublysodon (A. amplus and A. cristatus) based on unserrated
premaxillary teeth from the Lance Formation of Wyoming. These are juvenile tyrannosaurines,
based on the lack of serrations (Currie, 2003), and are thus probably Tyrannosaurus
rex, based on provenance. They are indistinguishable from Judith River tyrannosaurine
(Daspletosaurus?) juvenile premaxillary teeth, so are technically nomina
dubia. Hence neither species name can be a senior synonym of rex.
Manospondylus gigas- In 1892, Cope described Manospondylus
as a ceratopsid from the Lance Formation of South Dakota. Hatcher et al. (1907)
later referred it to the Theropoda, and Osborn (1916) noted its close resemblence
to Tyrannosaurus. While near certainly synonymous with T. rex,
as no other large theropods are known from Late Maastrichtian US deposits, the
holotype two cervicodorsal centra do not possess T. rex apomorphies other
than their size. Consequently, M. gigas has been viewed as invalid for
a century and is technically a nomen oblitum, so cannot have taxonomic priority
over T. rex despite its historical priority. New remains supposedly from
the Manospondylus holotype were discovered in 2000, as discussed below.
Armored Tyrannosaurus?- In 1900, the holotype of Dynamosaurus
imperiosus (then AMNH 5866) was discovered with 77 osteoderms (now NHMUK
R8001), thought by Osborn (1905, 1906, 1916) to belong to the theropod. Carpenter
(2004) confirmed these belong to Ankylosaurus, with the supposed differences
noted by Brown (1908) and Osborn being due to comparisons with AMNH Euoplocephalus
(or Anoplosaurus?) material.
The Nanotyrannus problem- Discovered in 1942 in the Hell Creek
Formation of Montana, CMN 7541 was described as Gorgosaurus lancensis
(Gilmore, 1946). It was generally assigned to this genus or its subjective synonym,
Albertosaurus (Russell, 1970; Paul, 1988), though Paul did place it in
a separate subgenus. In 1988, Bakker et al. redescribed the specimen as a new
genus, Nanotyrannus, and placed it as the most basal tyrannosauroid.
A bibliographic listing of the paper (in Currie, 1987) prior to its publication
used the name "Clevelanotyrannus", which was perhaps an early suggested
name for the taxon, though Currie (pers. comm. to Ford on www.paleofile.com)
claims he has never heard of it. Additionally, a New Scientist article (Anonymous,
1988) from right before the publication of Bakker et al.'s paper called it "Nanotyrannes".
Rozhdestvensky (1965) was the first to suggest CMN 7541 was a juvenile Tyrannosaurus,
which was also considered a possibility by Carpenter (1992), though Carr (1999)
was the first to officially propose it. Since then it has been clear that CMN
7541 is juvenile (due to striated cortical bone and numerous characters seen
in other juvenile tyrannosaurids), but it is disputed whether it is a juvenile
Tyrannosaurus rex (Holtz, 2001; Carr and Williamson, 2004; Carr, 2005;
Henderson, 2005), or the juvenile of a sister species to T. rex (Currie,
2003; Currie et al., 2003; Witmer and Ridgely, 2010; Larson, 2013). In 2001,
an additional juvenile specimen (BMRP 2002.4.1 or "Jane") conspecific
with CMN 7541 was discovered in the Hell Creek Formation of Montana. It was
discussed extensively at 2005 conferences, and is being monographed by Carr
et al.. Evidence for CMN 7541 and BMRP 2002.4.1 being distinct from T. rex
include a higher tooth count, subnarial foramen enclosed by maxilla, dorsally
opening jugal foramen, notches in the dorsal quadratojugal, lateral pneumatic
foramen on the quadratojugal, low cultriform process, small subsellar recess,
laterally positioned vagus foramen, two pairs of foramina in the basisphenoid
sinus, extensive medial subcondylar recess, adjacent medial and lateral foramina
in the subcondylar recess, strong condylotuberal crest, laterally oriented scapular
glenoid, fused scapulocoracoid and pevic sutures, and posteriorly hooked preacetabular
process. However, no Hell Creek tyrannosaurine adults with these characters
are known, nor are any juveniles lacking them. Schmerge and Rothschild (2016)
present a highly flawed paper proposing the lateral dentary groove of Jane is
an albertosaurine character, but don't even reference juvenile Tarbosaurus
IGM 107/7 which has the groove despite at least several adults lacking it, and
only code Nanotyrannus for a minority of known characters in their reanalysis
of the Brusatte et al. tyrannosauroid matrix. I provisionally accept Nanotyrannus
as a juvenile Tyrannosaurus rex, though the publication of BMRP 2002.4.1's
description may change this.
Additional specimens referred to Nanotyrannus consist mostly of teeth,
and have not been described in detail. Langston (1955 field notes) apparently
noted a skull and skeleton (presumably referred to Gorgosaurus lancensis
at the time) from the Late Cretaceous of Alberta, though these have not been
discussed in the literature since. Three teeth (DMNH coll.) from the Lance Formation
of South Dakota were referred by Bakker et al. (1988). Derstler (1995) reported
teeth from the Lance Formation of Wyoming. Another three teeth and a jugal (BHI
coll.) from the Hell Creek Formation of South Dakota associated with FMNH PR2081
were originally identified a s a juvenile T. rex (Erickson, 1999), but
have been referred to Nanotyrannus as well (Larson pers. comm., 1997
to Ford and Chure, 2001). A lacrimal (BHI 6235?) may also belong to this specimen
(Larson, 2013). It was reported that Nanotyrannus teeth (as identified
by Bakker) were associated with the subadult T. rex nicknamed Tinker
from the Hell Creek Formation of South Dakota (Blasing, DML 2006), though its
teeth are similar to those of adult tyrannosaurids. Spencer et al. (2001) referred
teeth from the Lance Formation of Wyoming to Nanotyrannus sp.. Kemmick
(2004) reported fifty Nanotyrannus teeth associated with what was then
thought to be a T. rex skeleton in Montana. This turned out to be the
skeleton of a different species from the earlier Judith River Formation however,
and these teeth are more likely from another juvenile tyrannosaurid. Maltese
(pers. comm., 2008) found these teeth were similar to albertosaurines and Daspletosaurus
in morphology. It should be noted that Nanotyrannus teeth only differ
from T. rex in ontogenetic characters, so isolated teeth cannot be referred
to either taxon. A phalanx was reported by Stenerson and O'Conner (1994) from
the Hell Creek Formation of South Dakota, but this is obviously based on size
alone. Larson et al. (2003) note that in their collection of ninety-one tyrannosaurid
teeth from the Lower Hell Creek Formation of South Dakota, some are more laterally
compressed than others, and that this includes large teeth, while small teeth
can be robust as well. They suggested the possibility of two tyrannosaurid taxa.
A largely complete specimen from the Hell Creek Formation of Montana associated
with a Triceratops (the so-called Dueling Dinosaurs) has been referred
to Nanotyrannus as well, but is in a private collection and remains undescribed.
It shares the high tooth count, enclosed subnarial foramen, jugal foramen orientation,
and hooked preacetabular process with other specimens, and additionally differs
from adult T. rex in having a phalanx on manual digit III.
Huxley rex- First observed in 1946, RTMP 81.12.1 (nicknamed Huxley rex)
is known from a badly eroded skeleton in the Scollard Formation of Alberta.
Only a pedal phalanx had been collected as of 1970 (Langston, 1965; Russell,
1970), though more was collected by Currie in 1981.
Dinotyrannus megagracilis- In 1967 a partial skeleton (LACM 23845)
was discovered in the Hell Creek Formation of Montana and initially thought
to be an immature Tyrannosaurus rex. It was described by Molnar (1980)
as an individual of Albertosaurus lancensis, now agreed to be a juvenile
T. rex or the juvenile of a sister species to T. rex. LACM 23845
was later (Paul, 1988) made the holotype of a new species- Albertosaurus
"megagracilis". Olshevsky (1995) placed the species in a new genus,
Dinotyrannus, which he believed was a derived tyrannosaurine closely
related to Nanotyrannus and Tyrannosaurus. Later, Rauhut (2000)
noted Albertosaurus "megagracilis" is a nomen nudum, as Paul
did not illustrate it, cites the wrong reference and gives no formal diagnosis.
This makes Olshevsky the official author of the taxon. Carr and Williamson (2000)
provisionally considered Dinotyrannus a subadult T. rex, which
confirmed in the detailed redescription and analysis by Carr and Williamson
(2004). The latter authors also corrected some misidentifications by Molnar,
such as the apparently downbent nasals being damaged, the supposedly absent
olecranon process of the ulna being missing, and the supposed manual ungual
being pedal ungual I. Carr and Williamson's identification is universally accepted
today.
The largest skull- Though discovered in 1967 and described in the technical
literature (Molnar, 1991), MOR 008 was not well known to the public until 2006,
when the incomplete skull was assembled and discovered to be larger than that
of FMNH PR2801. This makes the specimen, from the Hell Creek Formation of Montana,
the largest fairly complete Tyrannosaurus skull known.
Texas maxilla- In 1970, a maxilla was discovered in the Tornillo Formation
of Texas, described in Lawson's (1972) unpublished thesis as Tyrannosaurus
"vannus" (while names occuring only in theses are generally excluded
from this website, it was mentioned in the literature by Naish, 2009). It was
later described by Lawson (1976) as merely a subadult Tyrannosaurus rex.
Carpenter (1990) questioned this on the basis of the shorter anterior body,
deeper posteroventral process and slightly larger maxillary fenestra. However,
Carr and Williamson (2000) noted it shares numerous T. rex apomorphies
and that short anterior bodies are present in some other T. rex specimens
(e.g. BHI 3033). The proportional differences can thus be explained by individual
variation. Molnar (1991) and Brochu (2003) also accept this specimen as T.
rex or a sister species.
The largest maxilla- Collected in 1977, UCMP 118742 is a very large maxilla
(810 mm long) from the Hell Creek Formation of Montana. It is famous due to
Paul's (1988) estimate of a body length of 13.6 meters, which would make it
one of the longest Tyrannosaurus' known. In 1996 however, Paul (DML)
had stated his prior mass estimate (12 tons) was too high. His new mass estimate
(7-8.5 tons) is still 15% larger than his estimate for FMNH PR2801, so UCMP
118742 may still be 5% longer than FMNH PR2801 in his view, at ~13.4 meters.
Thus it seems Paul was revising his mass estimates of Tyrannosaurus,
not his length estimate of UCMP 118742.
The Jordan theropod or Stygovenator molnari- Molnar (1978) described
a partial theropod snout (LACM 28471) discovered in 1966, from the Hell Creek
Formation of Montana. He did not name it (calling it the Jordan theropod) and
identified the specimen as a dromaeosaurid. Currie (1987) suggested it may be
referrable to Aublysodon, and Paul (1988) later named it Aublysodon
molnaris (later emmended to molnari by Paul in 1990, to match the
gender of Aublysodon). Molnar and Carpenter (1989) redescribed the specimen
as Aublysodon cf. mirandus, due to the lack of difference between it
and the holotype tooth of that species. Olshevsky (1995) separated LACM from
Aublysodon as Stygivenator molnari, based on the supposedly smaller
and mesiodistally narrower premaxillary tooth than that of A. mirandus.
Carr and Williamson (2000) noted the supposedly diagnostic characters were typical
of juvenile tyrannosaurids and considered it the juvenile of an indeterminate
tyrannosaurid, pending restudy. Holtz (2001) included it in a cladistic analysis,
where it emerged as a basal tyrannosauroid along with a chimaera of Alectrosaurus
+ GI 100/50 + 100/51 and OMNH 10131 (a juvenile Bistahieversor) in an
"aublysodontine" clade. Currie (2003) considered LACM 28471 to be
a juvenile Tyrannosaurus rex, which was confirmed in a detailed redescription
and analysis by Carr and Williamson (2004). The latter authors also corrected
some misidentifications by Molnar, Molnar and Carpenter, and Olshevsky, such
as the presence of interdental plates and the identification of the supposed
premaxillary tooth as a first maxillary tooth. Most authors agree with the synonymy
with T. rex (including Holtz, 2004), with Olshevsky being an exception.
If Nanotyrannus lancensis turns out to be distinct from T. rex,
it is unclear which taxon the younger LACM 28471 belongs to.
Black Beauty and Stan- Discovered in 1980, RTMP 81.6.1 (nicknamed Black
Beauty) was discovered in the Willow Creek Formation of Alberta.
BHI 3033 (nicknamed Stan) was discovered in the Hell Creek Formation of South
Dakota in 1987 and excavated in 1992. It is exceptionally complete, especially
the skull (missing only one coronoid and articular) and vertebral column (missing
only less than fifteen caudals), though suffering numerous pathologies. The
specimen has been fully prepared but only the skull has been described (Larson,
2008). A portion of the BHI's website is devoted to the specimen- http://www.bhigr.com/pages/info/info_stan.htm.
Wankel Rex or T. regina-
Discovered in 1988, Wankel Rex was found in the Hell Creek Formation of
Montana. Originally catalogued as MOR 555, it was the subject of
Horner and Lessem's (1993) book "The Complete T. rex", and
has since been loaned to the USNM in June 2013 for 50 years (Freedom du
Lac, 2013) and is now known as USNM 555000. Paul et al. (2021)
have used this as the holotype of their proposed gracile Tyrannosaurus species, T. regina.
Their species diagnosis is- "Generally gracile with an adult
femur-length/circumference ratio over 2.4, usually one slender anterior
incisiform dentary tooth."
Sue or T. "stanwinstonorum" or T. imperator- Perhaps the most famous
Tyrannosaurus specimen, FMNH PR2081 (nicknamed Sue) was discovered in
1990 in the Hell Creek Formation of South Dakota. FMNH PR2081 is significant
for both its size (~12.8 m) and completeness. After a legal battle over who
owned the specimen, it was sold to the FMNH for $8.4 million. This is the most
complete specimen to be well described in the literature, with an extensive
osteology published (Brochu, 2003). A possible proatlas arch is preserved, the
first identified in a theropod. The furcula identified by Brochu and mounted
on the skeleton is a pathological gastralium (Larson and Rigby, 2005). However,
the latter authors identified the supposed thirteenth dorsal rib described by
Brochu as the true furcula. The supposed huge olfactory bulbs are actually olfactory
chambers, containing nasal turbinates (Witmer and Ridgely, 2005). Also notable
is that the remains of three other younger Tyrannosaurus were found with
the specimen (Larson, 1995), perhaps indicating social behavior. These have
not been described, however. Harlan (1990) used the name Tyrannosaurus gigantus
in the title of a Newsweek article about Sue, but did not intend this as an
actual species. Pickering (1995) made BHI 2033 (which FMNH PR2081 was catalogued
as until 2000) the holotype of a new species, Tyrannosaurus "stanwinstonorum".
This was published in a private newsletter however, so is a nomen nudum. It
was also based on characters which are probably individual variation (larger
body size than T. rex; reduced nasal rugosities), incorrect (palatine
recess absent; rugosity absent on ventral pterygoid wing of palatine; supradentary
absent), or ambiguous (reduced postorbital-orbital joint). There is therefore
no evidence T. "stanwinstonorum" is valid. Paul et al. (2021) have since used this as a holotype of their proposed early and robust Tyrannosaurus species T. imperator,
with the diagnosis of- "Generally robust with an adult
femur-length/circumference ratio of 2.4 or less; usually two slender
anterior incisiform dentary teeth." FMNH PR2081 has a website
devoted to it- https://www.fieldmuseum.org/blog/sue-t-rex.
Early 90's specimens- Discovered in 1991, SMNH P2523.8 (nicknamed Scotty)
is represented by an incomplete skull and skeleton from the Frenchman Formation
of Saskatchewan (Tokaryk and Bryant, 2004). The skeleton's size and arrangement,
and the composition of the surrounding matrix, have delayed preparation and
description, but the skull is being described by Tokaryk.
A specimen nicknamed Samson was excavated in 1992 in the Hell Creek
Formation of South Dakota. It originally went by the nicknames Z-rex
and Mr. Zed while it was for sale in Kansas. The CMN acquired it and
begain preparation of the exceptionally well preserved skull in 2004
and completed it in 2006. It was sold to a private collection in
2009. Glut (2002) reported the femur is 1.36 meters long, but
Larson (2008b) has it as 1.295 meters.
A specimen nicknamed Bowman was discovered in 1992 in the Hell Creek Formation
of North Dakota, briefly mentioned by Oakland and Pearson (1995). It is still
encased in plaster jackets and may not be prepared due to the hard concretion
surrounding the bones.
Discovered in 1993 is BHI 4100 (nicknamed Duffy), from the Hell Creek Formation
of South Dakota (Browne, 1993).
Discovered in 1994, BHI 4182 (nicknamed Fox or County rex) is based on a fragmentary
skull and skeleton from the Hell Creek Formation of South Dakota. Its dentary
is 90% as long as FMNH PR2081.
A specimen nicknamed Barnum was collected from the Hell Creek Formation of South
Dakota in 1995. Although it was popularized as being the rest of the Dynamosaurus
type specimen, both specimens preserve dentaries and a left femur, so this cannot
be the case (Ford, vrtpaleo; Carpenter, DML 2004). Unfortunately, it was sold
to a private bidder in an auction in 2004.
Discovered in 1995, LDP 977-2 (nicknamed Pete) was found in the Lance Formation
of Wyoming. Derstler and Myers (2008) wrote a preliminary report on it.
Rigby rex or Peck's rex- MOR 980 (nicknamed the Rigby rex then Peck's
rex) was collected and first reported in 1997 from the Hell Creek Formation
of Montana. It was originally said to be the largest Tyrannosaurus known,
with a pubis reportedly 8% longer than in FMNH PR2081. It was also said to have
larger, more robust forelimbs than T. rex and different caudal structure.
The pubis seemed too large for the cranial material, intitially suggesting different
proportions than other T. rex specimens. These differences caused Melbourne
(1998) to suggest some were calling the specimen Tyrannosaurus “imperator”,
though this is a nomen nudum and none of the differences have been substantiated
after further preparation. Later, Rigby claimed at least one other individual
was represented (as shown by the supposed presence of four pubes in the collection),
which was supposedly average sized. Another more fragmentary specimen was also
said to be possibly present. However, further preparation has confirmed the
presence of only one specimen in the quarry (Morrow pers. comm., 2006; Derstler
and Myers, 2008). At ~12.8 meters, it is indeed one of the largest T. rex
specimens and also one of the most complete (80%+). MOR 980 is also notable
for preserving a furcula (Larson and Rigby, 2005) and the first reported Tyrannosaurus
metacarpal III. A website was devoted to the specimen from 2004-2006- https://web.archive.org/web/20060808040539/http://www.pecksrex.com/.
Bucky- CMI 2001.90.1 (nicknamed Bucky) was discovered in 1998 in the
Hell Creek Formation of South Dakota. It is a subadult specimen notable for
its furcula (Larson and Rigby, 2005), the first correctly identified Tyrannosaurus
furcula to be described. The rest of the specimen remains undescribed, but is
featured on the BHI website (BHI 2007 online).
Alaskan Tyrannosaurus?- Gangloff (1998) listed Tyrannosaurus
sp.(?) in the faunal list for Alaskan dinosaurs, and only the Prince Creek
Formation is young enough to contain the genus. However, in a later work detailing
the theropod teeth from the Prince Creek Formation (Fiorillo and Gangloff, 2000),
the nine tyrannosaurid teeth were not identified to genus level. It is assumed
Gangloff reconsidered his tentative identification and there remains no Tyrannosaurus
known from Alaska.
Tinker the subadult- In 1998, a subadult Tyrannosaurus was discovered
in the Hell Creek Formation of South Dakota and nicknamed Tinker. Although touted
as a juvenile in the press releases, Tinker is much larger than the 'Nanotyrannus'
specimens CMNH 7541 and BMRP 2002.4.1, almost the size of the Dinotyrannus
holotype. It is therefore unsurprising it possesses a low number of mediolaterally
thick teeth characteristic of older tyrannosaurids, instead of the narrower
more numerous teeth of 'Nanotyrannus' specimens. Interestingly, the latter
type of tooth was found associated with Tinker, perhaps suggesting scavenging
by younger Tyrannosaurus individuals or social behavior. Blasing (DML
2006) stated that another young Tyrannosaurus (nicknamed Belle) and remains
of an adult (nicknamed Regina) were present in the jackets with Tinker. Unfortunately,
Tinker was not deposited in a museum and was bought by a private individual
who currently has it and Regina on display and for sale at the Etihad Modern
Art Gallery.
Manospondylus redescovered?- Disvovered in 1999 is BHI 6248 (nicknamed
E.D. Cope). These remains were found in the Hell Creek Formation of South Dakota,
possibly at the site Manospondylus' holotype was excavated from (based
on centra piled up at the site). This led Larson to propose it could be from
the same individual. Although Larson (in Anonymous, 2000a) suggested this could
make Manospondylus the valid name for Tyrannosaurus, this could
not happen. The fourth edition of the ICZN dictates that Manospondylus,
having been considered invalid for fifty years, is a nomen oblitum which cannot
replace a valid name such as Tyrannosaurus.
Horner's 2000 Hell Creek Project- Discovered in 2000 in the Lower Hell
Creek Formation of Montana, MOR 1125 (nicknamed B-rex) became famous in 2005
when Schweitzer et al. described medullary bone from its hindlimb elements.
This tissue is unique to female birds among extant animals and indicates the
specimen was a female as well. It is also unique among described Tyrannosaurus
specimens in being from the lower part of the Hell Creek Formation (Early Maastrichtian),
as opposed to others which are from the Late Maastrichtian. Of course with so
many undescribed specimens known, and so many specimens collected by amatuers,
it's possible other known Hell Creek Tyrannosaurus' are equally old.
For instance, the MOR website gives MOR 1131 the same locality number as MOR
1125, and notes MOR 1126 and 1128 are also from the Lower Hell Creek Formation.
Another famous T. rex specimen was found in 2000, MOR 1126 (nicknamed
C-rex or Celeste). Discovered in the Lower Hell Creek Formation of Montana,
this specimen is said to be ten percent larger than FMNH PR2081 (Anonymous,
2000b) and have a tibiofemoral ratio of 1.0. However, Larson (2008a) lists neither
femur nor tibia in the known material. At 14 meters, this would be one of the
largest Tyrannosaurus yet discovered, but this must be regarded as tentative
until the remains are prepared.
Additional specimens discovered in the same field expedition as MOR 1125 and
1126 include MOR 1127 (nicknamed L-rex), MOR 1128 (nicknamed G-rex), MOR 1131
(nicknamed J-rex) and MOR 1142 (nicknamed X-rex). MOR 1142 was originally thought
to be a Tyrannosaurus, but turned out to be an Edmontosaurus,
hence its nickname.
Post-2000 discoveries- MOR 1152 (nicknamed Frank or F-rex) is an additional
specimen known from the Lower Hell Creek Formation of Montana. It was discovered
in 2001.
The USNM are preparing a specimen found in 2001, in the Hell Creek Formation
of Montana. It has been nicknamed Nathan or N-rex.
In 2002, BHI 6230 (nicknamed Wyrex and later transferred to become HMNS 2006.1743.01)
was discovered in the Hell Creek Formation of Montana. This fairly complete
specimen is notable for preserving third metacarpal, the first radiale known
from a Tyrannosaurus, and the first skin impressions from the genus (Larson,
2008a). The impressions appear to be scaly and were described in detail by Bell
et al. (2017). As of 2004, many bones had been prepared. A website containing
numerous photographs of the specimen can be seen here- https://web.archive.org/web/20210816160204/http://www.unearthingtrex.com/.
Also in 2002, a specimen being prepared in the RMDRC (=AMNH 30564) (nicknamed
Sir William) was discovered in Montana. Originally identified as a T. rex
(Erickson et al., 2002; Kemmick, 2004), the specimen was reidentified as a new
taxon close to the ancestry of T. rex by Stein and Triebold (2005).
How big was T. rex and which specimen is largest? There have been
several contenders for the title of largest Tyrannosaurus- MOR 008, UCMP
118742, FMNH 2081 (Sue), MOR 980 (Rigby rex or Peck's rex) and MOR 1126 (Celeste
or C-rex). Only FMNH 2081 is known from a fairly complete skeleton, and only
it has been extensively described and illustrated in the technical literature
(although MOR 008 and UCMP 118742 have both been mentioned in reviews of Tyrannosaurus
morphology- e.g. Molnar, 1991; Currie, 2003; Carr, 2005). The mounted skeleton
of FMNH 2081 is 12.8 meters long, and less complete specimens are scaled to
it on this website. MOR 008's skull is stated to be 1.5 m, compared to FMNH
2081's 1.394 m. If the skeleton were in proportion, it would be 13.8 meters
long. However, the maxilla is only 84% as long, with a toothrow 90% as long.
The dentary is 87% as long with a toothrow 90% as long. These measurements suggest
a total length of 10.8-11.5 meters. UCMP 118742's maxilla was said to be 29%
longer than AMNH 5027 by Paul (1988), but is actually only 14% longer, with
a toothrow 18% longer (Larson, 2008b). If the skeleton were in proportion to
FMNH PR2081 (which has a 861 mm long maxilla and 645 mm toothrow), it would
be 12.1-12.4 meters long. MOR 980's mounted skeleton is said to be 12.8 meters
long, although its pubis was reportedly 8% longer than FMNH PR2081's. The skull
as reconstructed for sale on its website is slightly smaller than FMNH PR2081.
Finally, no measurements have been made for MOR 1126, merely Horner's estimate
that it is 10% longer than FMNH PR2801, which would make it 14.1 meters. One
point which needs to be made is that Tyrannosaurus individuals did not
all have the same proportions. For instance, FMNH PR2081's maxilla is 25% longer
than the holotype's. The scapula is 20% longer, the dentary 15% longer, metatarsal
IV 4% longer, the femur 3% longer, the sacrum 1% longer, the tibiae are equal
in length, and metatarsal II is actually 5% shorter. This brings some perspective
to the potentially confusing MOR 980 measurements noted above. It also suggests
caution when estimating the total length of fragmentary individuals. If only
FMNH PR2081's maxilla were known, we might suggest it was 25% larger than the
holotype, or 15.5 meters! Yet it was <5% larger, as the skeleton shows. So
maybe MOR 008 and UCMP 118742 had smaller bodies than their cranial remains
would suggest as well. As for MOR 1126, Horner's guess has little value until
measurements are taken.
Tyrannosaurus defined- Holtz (2001) defined Tyrannosauridae as
all taxa closer to Tyrannosaurus than to Aublysodon, as he advocated
a basal group of tyrannosauroids (aublysodontids) containing LACM 28471 (which
he assigned to Aublysodon), OMNH 10131 and Alectrosaurus (a chimaera
as used by Holtz). Tyrannosauridae would then contain the taxa closer to Tyrannosaurus
than to this clade- Gorgosaurus, Albertosaurus, Daspletosaurus,
Alioramus, Shanshanosaurus, Tarbosaurus and Tyrannosaurus
itself. However, LACM 28471 turned out to be a juvenile T. rex, OMNH
10131 a juvenile Bistahieversor, and the Aublysodon's holotype
(which Phylocode dictates the definition be based on) is indeterminate. It is
most likely a juvenile tyrannosaurine and may be Daspletosaurus based
on its age (see entry). The discovery of an apparent possible ancestor of T.
rex by Stein and Triebold (2005) in the same formation as Aublysodon's
holotype means it may even belong to T. rex's sister species. Thus, of
all tyrannosauroids, only T. rex specimens themselves can be confirmed
to be more closely related to the T. rex holotype than to the Aublysodon
holotype. This makes Holtz's definition of Tyrannosauridae synonymous in known
content to T. rex.
A similar situation occurs with Sereno's (1998) definition of Tyrannosauridae,
which was all taxa closer to Tyrannosaurus than to Aublysodon,
Alectrosaurus or Nanotyrannus (the latter three again being 'aublysodontids'
in Sereno's view). This case is more explicit though, as Nanotyrannus
is currently believed to be a juvenile T. rex or a juvenile of its sister
species. So at best Sereno's Tyrannosauridae encompasses only T. rex
itself, and at worst it encompasses some unidentified population of T. rex
individuals more closely related to CMN 9380 than to CMN 7541.
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