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)
= 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.
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.
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.
Flower, 1929. Reptilia and Amphibia. In Sclater (ed.). Records of zoological literature relating chiefly to the year 1928. The Zoological Record, London. 45, 1-77.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
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.
Kurzanov, 1989. O proiskhozhdenii i evolyutsii infraotryada dinozavrov Carnosauria. Paleontologicheskiy Zhurnal. 1989(4), 3-14.
Bonaparte, Novas and Coria, 1990. Carnotaurus sastrei Bonaparte, the horned, lightly built carnosaur from the Middle Cretaceous of Patagonia. Natural History Museum of Los Angeles County Contributions in Science. 416, 41 pp.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 169-209.
Sereno and Arcucci, 1990. The monophyly of crurotarsal archosaurs and the origin of bird and crocodile ankle joints. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 180(1), 21-52.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Sereno, Wilson, Larsson, Dutheil and Sues, 1994. Early Cretaceous dinosaurs from the Sahara. Science. 266, 267-271.
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.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku Saizensen. 9, 92-119 (part 1); 10, 75-99 (part 2).
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, 1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation. Science. 272(5264), 986-991.
Pharris, DML 1997. https://web.archive.org/web/20201115172819/http://dml.cmnh.org/1997Jun/msg00271.html
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen. 210(1), 41-83.
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). Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. Museum National d'Histoire Naturelle Laboratoire de Paleontologie. 329 pp.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmólska (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 Società 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, 70.
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. Cretaceous Research. 51, 35-55.
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

Tyrannosauroidea incertae sedis

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 pantyrannosaurian 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%).
Is Aviatyrannis an ornithomimosaur? Hattori et al. (2023) suggested "Because the ilium of Tyrannomimus is strikingly similar to the holotypic ilium of Aviatyrannis ..., its purported affinity to a tyrannosauroid is tested here against an alternative hypothesis that it in fact belongs an ornithomimosaur." Examination of the matrix shows Aviatyrannis was scored as different than other tyrannosauroids in- character 425-0 ("Ventral_edge_of_anterior_ala_of_ilium_ straight_or_gently_curved"), while it is actually hooked more than e.g. Juratyrant and the un-included Stokesosaurus; 435-1 ("Brevis_fossa_shape deeply_concave,_expanded_posteriorly_with_lateral_overhang"), which is also true in Stokesosaurus and probably Juratyrant, although the latter is crushed so that any posterior expansion was lost. Finally, Hattori et al. say "Although the dorsal half of the anterior margin is concave in Aviatyrannis, it is much shallower than the notch seen in tyrannosauroids. In addition, while some ornithomimosaurs lack the concavity on the anterior margin, many ornithomimosaurs do not have the corresponding portions preserved, so the conditions of these species remain unknown", but the presence of a concavity is still a character and the fact some ornithomimosaurs can't be scored in no way argues they might have the tyrannosauroid condition as well. Thus misscorings led to the placement of Aviatyrannis sister to Tyrannomimus in Choiniere's coelurosaur matrix as recovered by Hattori et al., and adding it (ilium only) and Tyrannomimus to Hartman et al.'s maniraptoromorph matrix results in a tyrannosauroid placement with an ornithomimosaur placement being two steps longer.
References- Zinke, 1998. Small theropod teeth from the Upper Jurassic coal mine of Guimarota (Portugal). Paläontologische 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.
Hattori, Shibata, Kawabe, Imai, Nishi and Azuma, 2023. New theropod dinosaur from the Lower Cretaceous of Japan provides critical implications for the early evolution of ornithomimosaurs. Scientific Reports. 13:13842.

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.

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 Knötschke, 2009)
?(DFMMh/FV 790.5) (juvenile?) tooth (2.8x1.9x1.3 mm) (Lubbe, Richter and Knötschke, 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 Knötschke, 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). Paläontologische 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.

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 Osmólska (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
?...(AMNH 6592) (~3 m) tooth, partial cervical vertebra, distal ungual, proximal ulna, phalanges, partial pubes
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.

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 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

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 Osmólska (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 Kundrát 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 Kundrát, 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, Kundrát, Sues, Williamson and Xu, 2016. Brains before brawn: Neurosensory evolution in tyrannosauroid dinosaurs. Journal of Vertebrate Paleontology. Program and Abstracts, 106.
Kundrát, Xu, Hančová, 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. Grundzüge der Paläontologie (Paläozoologie). II. Abteilung. Vertebrata. Druck und Verlag von R. Oldenbourg, München. 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.

undescribed possible pantyrannosaurian (USNM online)
Late Aptian, Early Cretaceous
Dinosaur Park / Cherokee-Sanford Brick Clay Pit / Muirkirk Clay Pit USNM 41614, Arundel Formation, Prince George's County, Maryland, US
Material- (USNM 540720) two right dentary fragments (~10.6 mm dorsovent along anterior dentigerous portion)
Comments- Discovered in 2010, USNM 540720 is an interesting specimen, consisting of two dentary fragments including portions of eight alveoli. Unlike allosauroids or eudromaeosaurs the interdental plates are widely separated, so this is neither a juvenile Acrocanthosaurus or Deinonychus, while the large alveoli present posteriorly at the point where the Meckelian groove widens and shallows eliminates even a basal ornithomimosaur like the grandis material. Among other theropods expected in Early Cretaceous North America, basal megaraptorans (e.g. Siats) and troodontids should differ in having fused interdental plates, while alvarezsaurs, therizinosaurs and oviraptorosaurs have much smaller alveoli compared to dentary depth, and archaeopterygids and Early Cretaceous avialans are much too small. Among remaining taxa with unfused interdental plates and full rows of relatively large dentary teeth, Eotyrannus has much taller interdental plates that contact and a taller sub-Meckelian area, Coelurus has a taller sub-Meckelian area, more dorsally placed paradental groove and is concave ventrally, while Ornitholestes also differs in the latter two features; Compsognathus has a taller Meckelian groove and more dorsally placed paradental groove; and Fukuivenator has a taller sub-Meckelian area, more dorsally placed paradental groove and a much more dorsoventrally expanded dentary posteriorly. The three most similar taxa are Sinornithosaurus, Richardoestesia and Nuthetes, all of similar size and with ventrally placed paradental grooves. Sinornithosaurus differs in having its paradental plates mostly fused except for the first four or five, Richardoestesia has a deeper Meckelian groove, and both of these have a taller sub-Meckelian area. Nuthetes uniquely matches the Arundel specimen in the very low paradental groove, so it is tentatively assigned the same phylogenetic identification here, although that genus is itself very similar to Richardoestesia and telling basal tyrannosauroids from basal dromaeosaurids based on mandibular remains is difficult at the best of times. Notably, teeth similar to Richardoestesia have been described from the Arundel (USNM 497748, 497749) with basal lengths of 2-4 mm, which matches the ~4 mm long alveoli of USNM 540720.

"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. Cretaceous 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 Osmólska (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 Arqueológico-Paleontológico Salense (ed.). Actas de las III Jornadas Internacionales sobre Paleontología 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 Holtz, 2004
Definition- (Tyrannosaurus rex <- Eotyrannus lengi)

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.

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.

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.

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 Osmólska (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
Paratypes- ?(CCMGE 432/12457) incomplete dorsal vertebra (93 mm) (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 854/16) partial braincase (Brusatte, Averianov, Sues, Muir and Butler, 2016)
?(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)
Referred- ?(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 477/12457) distal caudal vertebra (Nessov, 1995)
?(CCMGE 485/12457) anterior lateral tooth (Nessov, 1995)
? 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. 113(13), 3447-3452.
Brusatte, Muir, Averianov, Balanoff, Bever, Carr, Kundrát, Sues, Williamson and Xu, 2016. Brains before brawn: Neurosensory evolution in tyrannosauroid dinosaurs. Journal of Vertebrate Paleontology. Program and Abstracts, 106.

Eutyrannosauria Delcourt and Nelson Grillo, 2018
Definition- (Dryptosaurus aquilunguis + Tyrannosaurus rex) (modified from Delcourt and Nelson Grillo, 2018)
Comments- While unnamed specimens from the Campanian and Maastrichtian are listed below as eutyrannosaurs, the presence of Alectrosaurus suggests some may be basal tyrannosauroids (although that genus' phylogenetic placement and the Iren Dabasu Formation's age are poorly constrained).
Reference- Delcourt and Nelson Grillo, 2018. Tyrannosauroids from the Southern Hemisphere: Implications for biogeography, evolution, and taxonomy. Palaeogeography, Palaeoclimatology, Palaeoecology. 511, 379-387.

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.

"Bistityrannus" Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review in Dalman, online 2024
?= "Alamotyrannus" Dalman and Lucas, in press in Dalman, 2013 in part
"B. anax" Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review in Dalman, online 2024
?= "Alamotyrannus brinkmani" Dalman and Lucas, in press in Dalman, 2013 in part
Etymologies- Bisti is Navajo for "large area of shale hills" after the Bisti/De-Na-Zin Wilderness Area where the specimen was probably found + Latin tyrannus "ruler", common suffix for tyrannosauroids. Greek anax "tribal chief or leader".
Alamo after the Ojo Alamo Formation where the material is from, in turn named for the Ojo Alamo trading post + Latin tyrannus "ruler", common suffix for tyrannosauroids. brinkmani probably from vertebrate paleontologist Donald B. Brinkman.
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation?, San Juan Basin, New Mexico, US
Material- ?(ACM 7975; intended syntype of "Alamotyrannus brinkmani") anterior right dentary (~142 mm deep) (Dalman, 2013)
Campanian-Maastrichtian, Late Cretaceous
San Juan Basin, New Mexico, US
?(PMU.R35; = PMU.R85 of Carr and Williamson, 2000 Appendix 1; = PMU.R1235 of Carr and Williamson, 2000 Fig. 9G-H and Appendix 1; intended syntype of "Alamotyrannus brinkmani"?) anterior right dentary (Sullivan and Williamson, 1997)
Comments- ACM 7975 was discovered in July or August 1924, and figured and briefly described by Dalman and Lucas (2016) as "Tyrannosauridae indeterminate" despite the authors listing two characters purportedly differentiating it from other tyrannosaurids. Namely, "the possession of two foramina intermandibularis oralis" and "the morphology of the anterior step of the lingual bar" which "is anterodorsally inclined and has a relatively short posterior surface (towards the mouth), whereas in other tyrannosaurids (e.g., Daspletosaurus, Tarbosaurus, and Tyrannosaurus) the posterior surface of the step is well-developed and nearly the same length as the dorsal surface."
Dalman (2013) stated "the many isolated but diagnostic tyrannosaurid skeletal fossil elements from the Naashoibito Member of the Ojo Alamo Formation (early Maastrichtian) of northwestern New Mexico (Sullivan et al. 2005; Jasinski et al. 2011; Dalman and Lucas, in press) provide evidence for the occurrence of a new taxon of a large tyrannosaurid" with the in press paper's bibliographic entry naming it "Alamotyrannus brinkmani." Stuchlik (pers. comm. to Dalman, 7-2018) informs me the intended holotype was two dentaries, presumedly including ACM 7975 that was mentioned in Dalman (2013) as "the new Ojo Alamo tyrannosaurid taxon ACM 7975." Dalman (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. A probable explanation is that Dalman and Lucas (2016) briefly described and figured ACM 7975 as a diagnostic tyrannosaurid under study by Dalman but noted that the previous referral to the Ojo Alamo Formation was due to its discoverer Loomis using an old, broader definition for the formation. Geographical and preservational data indicated instead that ACM 7975 was probably from the De-na-zin Member of the Kirtland Formation, so mixing it with the diagnostic Ojo Alamo elements noted by Dalman (2013) would make his "Alamotyrannus" concept a chimaera. Thus Dalman would want to describe the diagnostic Kirtland dentary and the diagnostic Ojo Alamo elements as different taxa, which seems to be the plan based on his 2024 online curriculum vitae.
This lists "Dalman, S.G., Jasinski, S.E., Lucas, S.G., Malinzak, D.E., Loewen, M.A., Fiorillo, A.R., Currie, P.J. 2024. Bistityrannus anax, a new tyrannosaurid from the Kirtland Formation (Upper Cretaceous) of northwestern New Mexico. Cretaceous Research (in review/in progress)" under Published Research despite being unpublished as of 9-21-2024. The genus and species are obviously invalid pending this publication as his online curriculum vitae is not "issued for the purpose of providing a public and permanent scientific record" (IZCN Article 8.1.1), "produced in an edition containing simultaneously obtainable copies by a method that assures 8.1.3.1. numerous identical and durable copies (see Article 8.4), or 8.1.3.2. widely accessible electronic copies with fixed content and layout" (Article 8.1.3), does not "state the date of publication in the work itself, and" is not "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" (Articles 8.5.2 and 8.5.3), the taxa are not "accompanied by a description or definition that states in words characters that are purported to differentiate the taxon" (Article 13.1) or "explicitly indicated as intentionally new" (Article 16.1).
Another unpublished entry on that page with identical authorship is "Denazinosaurus sicarius, a new tyrannosaurid from the Kirtland Formation (De-na-zin Member) Upper Cretaceous of New Mexico, USA", giving us two proposed new Kirtland tyrannosaurids with nothing distinguishing them in their publication titles besides "Denazinosaurus" definitely being from the De-na-zin Member. While either or neither of these could be intended for former "Alamotyrannus" dentary ACM 7975, it's here suggested the stratigraphic uncertainty behind that specimen's discovery makes De-Na-Zin unlikely to feature in the article title or genus name. Thus ACM 7975 is probably "Bistityrannus", while "Denazinosaurus" would be a different specimen with a more definite locality. Given most De-Na-Zin tyrannosaurid specimens are isolated teeth and postcrania (generally considered indeterminate in Tyrannosauridae), that ideally Dalman would want a specimen comparable to ACM 7975 to erect a new contemporaneous species distinct from it, and that he is describing other tyrannosaurids based on dentaries (see Fruitland Formation KU VP-96888), the obvious identity of "Denazinosaurus" would be classic dentary USNM V 8346 described by Gilmore in 1916 (see entry). Whether this logic proves true awaits either publication.
Interestingly, given Dalman originally intended to name "Alamotyrannus" based on two dentary syntypes including ACM 7975, it implies a second tyrannosaur dentary plausibly from the Ojo Alamo Formation. While NMMNH P-7199 is an obvious possibility as the only dentary I know of confirmed from that formation, it's so poorly preserved that it's never even been figured. I propose a more likely possibility is anterior dentary PMU.R35 whose locality can only be specified to San Juan County, and which seems to have the anterior step morphology described for ACM 7975- anterodorsally inclined with a short posterior surface (Sullivan and Williamson, 1997- Fig. 1D). As its uncertain stratigraphic placement makes it plausibly from the De-Na-Zin Member, this may be an additional specimen of "Bistityrannus". Discovered in 1921 or 1922, this was not published until Sullivan and Williamson's 1997 review of PMU specimens sent by Sternberg, where it is figured in lateral and medial views as "tyrannosaurid partial right dentary."
References- 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. 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.
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.
Dalman and Lucas, 2016. Frederic Brewster Loomis and the 1924 Amherst College paleontological expedition to the San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 74, 61-66.
Dalman, 2024 online. https://www.montana.edu/earthsciences/graduate-program/students/cv/Sebastian_Dalman.html
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.
Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review a. Denazinosaurus sicarius, a new tyrannosaurid from the Kirtland Formation (De-na-zin Member) Upper Cretaceous of New Mexico, USA. Acta Palaeontologica Polonica.
Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review b. Bistityrannus anax, a new tyrannosaurid from the Kirtland Formation (Upper Cretaceous) of northwestern New Mexico. Cretaceous Research.

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 Osmólska (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.

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.

"Denazinosaurus" Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review in Dalman, online 2024
"D. sicarius" Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review in Dalman, online 2024
Etymology- De-Na-Zin is Navajo for "cranes" and presumedly based on the De-na-zin Member of the Kirtland Formation which the taxon would be from + Greek sauros "reptile." Latin sicarius "assassin", as it's a probable hunter.
Late Campanian, Late Cretaceous
head of Hunter Wash / locality 60 of Bauer 1916, De-na-zin Member of Kirtland Formation, San Juan County, New Mexico, US
Material- ?(USNM V 8346) incomplete left dentary (tooth row length 354 mm) (Gilmore, 1916)
Comments- Discovered August 20, 1915, this was initially described by Gilmore (1916) as "Deinodon?" and figured in medial view. He noted "In the number of tooth sockets this jaw agrees with Dynamosaurus imperosus Osborn,^a [now Tyrannosaurus rex] but in the general form of the dentary, particularly the contour of the anterior end, it approaches Albertosaurus⁴ (Dryptosaurus) most nearly, but as the dentary of Albertosaurus has sockets for 15 teeth the presence of 13 in this individual would appear to show its distinctness." Gilmore further stated "It is quite possible that this dentary pertains to the genus Deinodon, but that can not be determined at this time because the dentary of that genus is unknown. The identification of this specimen must therefore await the discovery of additional material." Gilmore (1920) repeated most of this verbatim, listing it as "DEINODON, species." this time. In a later publication, Gilmore (1935) wrote "Comparison of this bone directly with a dentary of Gorgosaurus libratus Lambe from the Belly River of Canada now shows such close resemblances in size, shape, and other characteristics down to the smallest details as to leave little doubt of their being congeneric. Likewise, the number of alveoli (13) is in agreement with Lambe’s (1917) determination from a number of specimens that the dentary in this genus bears 13 or 14 teeth." Lucas et al. (1987) felt the dentary "almost certainly pertain[s] to Albertosaurus" sensu Russell 1970, equating to modern Albertosaurinae. Carr and Williamson (2000) figured the dentary in lateral, medial and dorsal views, and stated "Based on personal observation of this and other tyrannosaurid dentaries, this bone does not appear to be diagnostic at the specific or generic levels." They noted "There are actually 14 dental alveoli" and concluded "Because the alveolus count of USNM 8346 overlaps with that of A. libratus and T. rex and in light of the fact that tooth count is variable in these species, we consider it best to consider this specimen as indeterminate Tyrannosauridae." Indeed, 14 dentary alveoli are also present in some Albertosaurus sarcophagus (AMNH 5222, TMP 1986.064.0001), Daspletosaurus torosus (CMN 8506) and Tarbosaurus (IGM 107/7) specimens.
Dalman's 2024 online curriculum vitae lists "Dalman, S.G. Jasinski, S.E., Lucas, S.G., Malinzak, D.E., Loewen, M.A., Fiorillo, A.R. and Currie, P.J. 2024. Denazinosaurus sicarius, a new tyrannosaurid from the Kirtland Formation (De-na-zin Member) Upper Cretaceous of New Mexico, USA. Acta Palaeontologica Polonica (in review/in progress)" under Published Research despite being unpublished as of 9-21-2024. The genus and species are obviously invalid pending this publication as his online curriculum vitae is not "issued for the purpose of providing a public and permanent scientific record" (IZCN Article 8.1.1), "produced in an edition containing simultaneously obtainable copies by a method that assures 8.1.3.1. numerous identical and durable copies (see Article 8.4), or 8.1.3.2. widely accessible electronic copies with fixed content and layout" (Article 8.1.3), does not "state the date of publication in the work itself, and" is not "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" (Articles 8.5.2 and 8.5.3), the taxa are not "accompanied by a description or definition that states in words characters that are purported to differentiate the taxon" (Article 13.1) or "explicitly indicated as intentionally new" (Article 16.1).
Another unpublished entry on that page with identical authorship is "Bistityrannus anax, a new tyrannosaurid from the Kirtland Formation (Upper Cretaceous) of northwestern New Mexico", giving us two proposed new Kirtland tyrannosaurids with nothing distinguishing them in their publication titles besides "Denazinosaurus" definitely being from the De-na-zin Member. While either or neither of these could be intended for former "Alamotyrannus" dentary ACM 7975 (see "Bistityrannus" entry), it's here suggested the stratigraphic uncertainty behind that specimen's discovery makes De-Na-Zin unlikely to feature in the article title or genus name. Thus ACM 7975 is probably "Bistityrannus", while "Denazinosaurus" would be a different specimen with a more definite locality. Given most De-Na-Zin tyrannosaurid specimens are isolated teeth and postcrania (generally considered indeterminate in Tyrannosauridae), that ideally Dalman would want a specimen comparable to ACM 7975 to erect a new contemporaneous species distinct from it, and that he is describing other tyrannosaurids based on dentaries (see Fruitland Formation KU VP-96888), the obvious identity of "Denazinosaurus" would be classic dentary USNM V 8346. Whether this logic proves true awaits either publication.
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, Mateer, Hunt and O'Neill, 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.
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.
Dalman, 2024 online. https://www.montana.edu/earthsciences/graduate-program/students/cv/Sebastian_Dalman.html
Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review a. Denazinosaurus sicarius, a new tyrannosaurid from the Kirtland Formation (De-na-zin Member) Upper Cretaceous of New Mexico, USA. Acta Palaeontologica Polonica.
Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review b. Bistityrannus anax, a new tyrannosaurid from the Kirtland Formation (Upper Cretaceous) of northwestern New Mexico. Cretaceous Research.

"Erinotonax" Dalman, Jasinski, Loewen, Malinzak, Currie, Fiorillo and Lucas, in progress/review in Dalman, online 2024
"E. sabathi" Dalman, Jasinski, Loewen, Malinzak, Currie, Fiorillo and Lucas, in progress/review in Dalman, online 2024
Etymology- Perhaps Greek Eri "very" + Greek notos "south" + Greek ánax "king" (suggested by Creisler, pers. comm. 9-2024). sabathi after paleontologist Karol Sabath.
Late Campanian, Late Cretaceous
Ne-nah-ne-zad Member, Fruitland Formation, San Juan Basin, New Mexico, US
Material- ?(KU VP-96888) (~10 m) anterior left dentary (140 mm deep) (Dalman and Lucas, 2021)
Comments- Dalman and Lucas (2021) state "The specimen is being described by us in detail in another paper as belonging to a new genus and a new species" but contradictorally identify it as "Tyrannosauridae indet." They also say "When compared to the dentaries of other tyrannosaurids such as Albertosaurus, Bistahieversor, Daspletosaurus, Gorgosaurus, Lythronax, Tarbosaurus, Teratophoneus and Tyrannosaurus, the body length of KUVP-96888 was close to 10 m." This is listed as Tyrannosaurus sp. in the KU online catalog.
Coincidentally, Dalman's 2024 online curriculum vitae lists "Dalman, S.G., Jasinski, S.E., Loewen, M.A., Malinzak, D.E., Currie, P.J., Fiorillo, A.R., Lucas, S.G. 2024. Erinotonax sabathi, a new tyrannosaurid from the Fruitland Formation (Upper Cretaceous) of New Mexico, USA, insights into the evolution, diversity, and paleogeography of tyrannosaurids in North America. PeerJ (in review/in progress)" despite being unpublished as of 9-21-2024. The genus and species are obviously invalid pending this publication as his online curriculum vitae is not "issued for the purpose of providing a public and permanent scientific record" (IZCN Article 8.1.1), "produced in an edition containing simultaneously obtainable copies by a method that assures 8.1.3.1. numerous identical and durable copies (see Article 8.4), or 8.1.3.2. widely accessible electronic copies with fixed content and layout" (Article 8.1.3), does not "state the date of publication in the work itself, and" is not "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" (Articles 8.5.2 and 8.5.3), the taxa are not "accompanied by a description or definition that states in words characters that are purported to differentiate the taxon" (Article 13.1) or "explicitly indicated as intentionally new" (Article 16.1).
Unless Dalman and Lucas are describing multiple new genera of tyrannosaurids from the Fruitland Formation, it seems likely "Erinotonax" is KU VP-96888. Given its fragmentary and pathological condition, I'm skeptical of any future diagnosis.
References- Dalman and Lucas, 2021. New evidence for cannibalism in tyrannosaurid dinosaurs from the Upper Cretaceous (Campanian/Maastrichtian) San Juan Basin of New Mexico. New Mexico Museum of Natural History and Science Bulletin. 82, 39-56.
Dalman, 2024 online. https://www.montana.edu/earthsciences/graduate-program/students/cv/Sebastian_Dalman.html
Dalman, Jasinski, Loewen, Malinzak, Currie, Fiorillo and Lucas, in progress/review. Erinotonax sabathi, a new tyrannosaurid from the Fruitland Formation (Upper Cretaceous) of New Mexico, USA, insights into the evolution, diversity, and paleogeography of tyrannosaurids in North America. PeerJ.

"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).

"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 nomen 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 Osmólska (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.

undescribed Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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.
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 Eutyrannosauria (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 Eutyrannosauria (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 wilsoni.
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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 eutyrannosaur (MOR online)
Late Maastrichtian, Late Cretaceous
Livingston Formation, Montana, US
Material- (MOR 002) postcranial skeleton (MOR online)

undescribed Eutyrannosauria (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 rex, 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 eutyrannosaur (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 eutyrannosaur (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 eutyrannosaur (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 Eutyrannosauria (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. These may belong to the contemporaneous Lythronax.
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 Eutyrannosauria (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 eutyrannosaur (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (Brown, 1910)
Late Campanian-Maastrichtian, Late Cretaceous
San Juan Basin, New Mexico, US
Material- (AMNH 2479 in part; syntype of Dysganus encaustus) two tooth fragments (Carr and Williamson, 2000)
(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.R36; = PMU.R56 of Carr and Williamson, 2000 Appendix 1) tooth (Sullivan and Williamson, 1997)
(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. 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 Eutyrannosauria (Carr and Williamson, 2000)
Late Campanian, Late Cretaceous
Fruitland Formation or Hunter Wash or Farmington Members of 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
Comments- These may belong to the contemporaneous Bistahieversor.
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 Eutyrannosauria (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)
Comments- These may belong to the contemporaneous Bistahieversor or "Erinotonax".
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 Eutyrannosauria (Carr and Williamson, 2000)
Late Campanian, Late Cretaceous
Hunter Wash or Farmington Members of Kirtland Formation, New Mexico, US
Material- (KUVP 12164) teeth
(NMMNH P-7178) partial phalanx
(NMMNH coll.) eight teeth [see Carr and Williamson, 2000 for numbers]
Comments- These may belong to the contemporaneous Bistahieversor.
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 Eutyrannosauria (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]
Comments- These may belong to the contemporaneous Bistahieversor.
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 Eutyrannosauria (Gilmore, 1916)
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-12999; = UNM FKK-080) tooth (Lucas et al., 1987)
(NMMNH P-13000; = UNM FKK-076) tooth (Lucas et al., 1987)
(NMMNH P-20879) right pedal phalanx IV-3 (Hunt and Lucas, 1992)
(NMMNH P-25071) distal femur (Carr and Williamson, 2000)
(NMMNH P-25083) (~7 m) right femur (810 mm) (Dalman and Lucas, 2021)
(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]
(UNM FKK-77; lost) tooth (Lucas et al., 1987)
(UNM FKK-78; lost) tooth (Lucas et al., 1987)
(UNM FKK-79; lost) tooth (Lucas et al., 1987)
(USNM 8355) (juvenile) premaxillary tooth (Gilmore, 1916)
Comments- Jasinski et al. (2011) state the teeth reported by Lucas et al. (1987) "were purportedly from the Naashoibito Member, but Carr and Williamson (2000) concluded that only NMMNH P-13000 was from that unit. They determined that the other teeth were from the stratigraphically lower De-na-zin Member (Kirtland Formation). Because all the specimens, including NMMNH P-13000, are from the same locality, we considered all of them as coming from the De-na-zin Member."
Some or all of this material may belong to the contemporaneous "Denazinosaurus" or "Bistityrannus".
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.
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.
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, 2021. New evidence for cannibalism in tyrannosaurid dinosaurs from the Upper Cretaceous (Campanian/Maastrichtian) San Juan Basin of New Mexico. New Mexico Museum of Natural History and Science Bulletin. 82, 39-56.

undescribed Eutyrannosauria (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 Eutyrannosauria (Lehman, 1985)
Late Maastrichtian, Late Cretaceous
Javelina Formation, Texas, US
Comments- These may belong to the contemporaneous Tyrannosaurus rex.
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 Eutyrannosauria (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.

unnamed Eutyrannosauria (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 eutyrannosaur (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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 Eutyrannosauria (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.

undescribed Eutyrannosauria (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 Eutyrannosauria (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.

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.

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 Osmólska (eds.). The Dinosauria Second Edition. University of California Press. 111-136.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (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.

"Atroxicarius" Dalman, Jasinski, Loewen, Lucas, Malinzak, Fiorillo and Currie, in progress/review in Dalman, online 2024
"A. eversor" Dalman, Jasinski, Loewen, Lucas, Malinzak, Fiorillo and Currie, in progress/review in Dalman, online 2024
Etymology- Latin atrox "fierce" + ? Latin arius "agent of use", except the 'c' is unexplained and atrox is not a noun. Creisler (pers. comm 9-2024) suggests atrox + Latin cara "head" + Latin suffix -ius, but that should imply cranial remains which are not at all certain. Greek eversor "destroyer."
Late Maastrichtian, Late Cretaceous
NMMNH L-3961, Naashoibito Member of Ojo Alamo Formation, San Juan County, New Mexico, US
?(NMMNH P-7199) partial left dentary, 113 tooth fragments, partial vertebra (Carr and Williamson, 2000)
Late Maastrichtian, Late Cretaceous
SMP 313b, Naashoibito Member of Ojo Alamo Formation, Hunter Wash, San Juan County, New Mexico, US
?(SMP VP-1848) incomplete left metatarsal I (~93 mm) (Jasinski, Sullivan and Lucas, 2011)
Late Maastrichtian, Late Cretaceous
SMP 371, Naashoibito Member of Ojo Alamo Formation, Betonnie Tsosie Wash, San Juan County, New Mexico, US
?(SMP VP-1113) incomplete right femur (~1 m) (Carr and Williamson, 2000)
Late Maastrichtian, Late Cretaceous
SMP 424b, Naashoibito Member of Ojo Alamo Formation, San Juan Basin, New Mexico, US
?(SMP VP-2105) incomplete right scapulocoracoid (coracoid ~253 mm proximodist) (Jasinski, Sullivan and Lucas, 2011)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, San Juan County, New Mexico, US
?(Ratkevich coll.) right metatarsal IV (~513 mm) (Lehman, 1981)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, San Juan Basin, New Mexico, US
?(AMNH 5882) right pedal phalanx IV-2 (~143 mm) (Carr and Williamson, 2000)
Comments- Discovered May 7, 1998, NMMNH P-7199 is a partial dentary associated with tooth fragments and a partial vertebra that all remain unfigured. Carr and Williamson (2000) refer it "to cf. T. rex on the basis of its large apical and midheight denticles, 7.5 denticles per 5 mm and 8.5 denticles per 5 mm, respectively" on the mesial carina and note the dentary and vertebra "are too weathered and incomplete to permit identification." Jasinski et al. (2011) suggested however that "identification below the family level (Tyrannosauridae) based on denticle densities is tenuous" so retained it as Tyrannosauridae indet.. The NMMNH online catalog incorrectly lists this as being from the De-na-zin Member of the Kirtland Formation.
Jasinski et al. (2011) notes "The scapulocoracoid (SMP VP-2105, Fig. 7A-B) may be from an adult as it compares readily in size and morphology to the scapulocoracoids of Tyrannosaurus rex (FMNH PR2081; Brochu, 2003, fig. 80). While this indicates the presence of a large tyrannosaurid in the Naashoibito Member, the specimen cannot be confidently referred to Tyrannosaurus rex, but may represent Tyrannosaurus sp."
Femur SMP VP-1113 is first published as "SMP VP-? femur Naashoibito Mbr., Kirtland Fm. Tyrannosauridae indet" by Carr and Williamson (2000), then figured by Lucas and Sullivan (2000) as "Tyrannosauridae, incomplete femur." Sullivan et al. (2005) identified it as "cf. Daspletosaurus sp., based on comparison with USNM 10754, a right femur of either Gorgosaurus or Daspletosaurus sp. (labelled as Albertosaurus sp.) ... from the Dinosaur Park Formation" because "the size and morphology of SMP VP-1113 closely resembles that of USNM 10754", but the only stated character is gracility compared to Tyrannosaurus, which is true for almost all other tyrannosaurids.
Jasinski et al. (2011) figure a metatarsal I (incorrectly called "the 1st phalanx of the 1st metatarsal" on page 226), which along with femur SMP VP-1113 they state is "from a large tyrannosaurid that seems more gracile than an adult Tyrannosaurus rex, although they may represent a juvenile or sub-adult."
Lehman (1981) wrote "single, complete, right fourth metatarsal (on loan from the collection of Ronald P. Ratkevich of Alamogordo, New Mexico) is tentatively referred here to Albertosaurus sp." using the broad concept of the genus popular at the time where it included Gorgosaurus. It is figured in anterior, medial and five sectional views.
Carr and Williamson (2000) reported "a large pedal phalanx (AMNH 5882; Fig. 4A-F) was collected from the Naashoibito Member of the Kirtland Formation. The collector and date of collection of this specimen are unknown" and figured it in multiple views as Tyrannosauridae indet.. Oddly, the AMNH online catalog lists this as being collected by Brown from the Hell Creek Formation of Snow Creek, Montana, but does correctly list it as a "2nd phalanx" of Theropoda. Williamson and Carr (2005) referred the phalanx to cf. Tyrannosaurus rex in an abstract.
"Alamotyrannus" into "Atroxicarius"- Dalman (2013) stated "the many isolated but diagnostic tyrannosaurid skeletal fossil elements from the Naashoibito Member of the Ojo Alamo Formation (early Maastrichtian) of northwestern New Mexico (Sullivan et al. 2005; Jasinski et al. 2011; Dalman and Lucas, in press) provide evidence for the occurrence of a new taxon of a large tyrannosaurid" with the in press paper's bibliographic entry naming it "Alamotyrannus brinkmani." Stuchlik (pers. comm. to Dalman, 7-2018) informs me the intended holotype was two dentaries, presumedly including ACM 7975 that was mentioned in Dalman (2013) as "the new Ojo Alamo tyrannosaurid taxon ACM 7975." Dalman (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. A probable explanation is that Dalman and Lucas (2016) briefly described and figured ACM 7975 as a diagnostic tyrannosaurid under study by Dalman but noted that the previous referral to the Ojo Alamo Formation was due to its discoverer Loomis using an old, broader definition for the formation. Geographical and preservational data indicated instead that ACM 7975 was probably from the De-na-zin Member of the Kirtland Formation, so mixing it with the diagnostic Ojo Alamo elements noted by Dalman (2013) would make his "Alamotyrannus" concept a chimaera. Thus Dalman would want to describe the diagnostic Kirtland dentary and the diagnostic Ojo Alamo elements as different taxa, which seems to be the plan based on his 2024 online curriculum vitae.
This lists "Dalman, S.G., Jasinski, S.E., Loewen, M.A., Lucas, S.G., Malinzak, D.E., Fiorillo, A.R., and Currie, P.J. 2024. Atroxicarius eversor, a new tyrannosaurid from the Ojo Alamo Formation (Upper Cretaceous) of New Mexico, USA, new insights into the evolution of bistahiversorin tyrannosaurids in North America. Anatomical Records (in review/in progress)" under Published Research despite being unpublished as of 9-21-2024 and presumedly referring to The Anatomical Record. This would then seem to be the new name of the Ojo Alamo tyrannosaurid, which the title suggests is related to Bistahieversor in a new tribe "Bistahieversorini", although note Dalman misspelled it without the first 'e'. The tribe, genus and species are obviously invalid pending this publication as his online curriculum vitae is not "issued for the purpose of providing a public and permanent scientific record" (IZCN Article 8.1.1), "produced in an edition containing simultaneously obtainable copies by a method that assures 8.1.3.1. numerous identical and durable copies (see Article 8.4), or 8.1.3.2. widely accessible electronic copies with fixed content and layout" (Article 8.1.3), does not "state the date of publication in the work itself, and" is not "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" (Articles 8.5.2 and 8.5.3), the taxa are not "accompanied by a description or definition that states in words characters that are purported to differentiate the taxon" (Article 13.1) or "explicitly indicated as intentionally new" (Article 16.1) and even a correctly spelled "bistahieversorine" is not in Latinized form (article 11.7.2) nor is it "accompanied by citation of the name of the type genus" (Article 16.2). Based on another Dalman et al. (in review/progress) citation in Dalman's curriculum vitae, the De-na-zin tyrannosaurid will be named "Denazinosaurus sicarius" and preseumedly includes dentary ACM 7975 as the intended holotype.
It's unknown which specimens will be types of "Atroxicarius", but as Dalman (2013) cited skeletal (not dental, which are generally undiagnostic) elements described by Sullivan et al. (2005) and Jasinski et al. (2011), they plausibly include dentary and vertebra NMMNH P-7199, scapulocoracoid SMP VP-2105, femur SMP VP-1113, metatarsal I SMP VP-1848, a metatarsal IV from the private Ratkevich collection, and/or pedal phalanx AMNH 5882.
References- 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.
Lucas and Sullivan, 2000. Stratigraphy and vertebrate biostratigraphy across the Cretaceous-Tertiary boundary, Betonnie Tsosie Wash, San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 95-103.
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.
Dalman and Lucas, 2016. Frederic Brewster Loomis and the 1924 Amherst College paleontological expedition to the San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 74, 61-66.
Dalman, 2024 online. https://www.montana.edu/earthsciences/graduate-program/students/cv/Sebastian_Dalman.html
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.
Dalman, Jasinski, Loewen, Lucas, Malinzak, Fiorillo and Currie, in progress/review. Atroxicarius eversor, a new tyrannosaurid from the Ojo Alamo Formation (Upper Cretaceous) of New Mexico, USA, new insights into the evolution of bistahiversorin tyrannosaurids in North America. The Anatomical Record.
Dalman, Jasinski, Lucas, Malinzak, Loewen, Fiorillo and Currie, in progress/review. Denazinosaurus sicarius, a new tyrannosaurid from the Kirtland Formation (De-na-zin Member) Upper Cretaceous of New Mexico, USA. Acta Palaeontologica Polonica.

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 (Archer and Babiarz, 1992)
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) (Lehman and Carpenter, 1990)
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.
Archer and Babiarz, 1992. Another tyrannosaurid dinosaur from the Cretaceous of northwest New Mexico. Journal of Paleontology. 66, 690-691.
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, Kundrát, 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.

Labocania Molnar, 1974
L. anomala Molnar, 1974
Late Campanian, Late Cretaceous
El Gallo Formation, Mexico
Diagnosis- pneumatic quadrate; oblique ridge on posterior face of quadrate; lateral dentary shelf.
Holotype- (IGM 5307; = LACM 20877) (~7.5 m; ~1.2 tons) (skull ~660 mm) partial maxilla, distal quadrate, frontal, dentary fragment, premaxillary teeth, maxillary teeth, chevron, ilial fragment, proximal ischium (~750 mm), shaft of pubis, nearly complete metatarsal II (<507 mm), phalanx III-2 (~115 mm), long bone fragments
Comments- Discovered in 1970, this taxon was originally compared to Indosaurus, Shaochilong and a possible metriacanthosaurid dentary from the Kalaza Formation (IVPP V903). Similarities to tyrannosaurids were also noted. Paul (1988) referred it to his Allosauridae, which was paraphyletic to tyrannosaurids. Lamanna and Smith reexamined this species in 1998 and found the specimen to be more fragmentary than expected. They conclude it is a tyrannosaurid, although it may not be valid. Chure (2000) referred it to Holtz's tyrannosaur + ornithomimosaur + troodontid clade, and thought it was most closely related to Shaochilong.
References- Molnar, 1974. A distinctive theropod dinosaur from the Upper Cretaceous of Baja California (Mexico). Journal of Paleontology. 48(5), 1009-1017.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 464 pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod family Allosauridae. PhD thesis. Columbia University. 964 pp.
Rivera-Sylva and Carpenter, 2014. Mexican saurischian dinosaurs. In Rivera-Sylva, Carpenter and Frey (eds.). Dinosaurs and Other Reptiles from the Mesozoic of Mexico. Indiana University Press. 143-155.
Ramírez-Velasco and Hernández-Rivera, 2015. Diversity of Late Cretaceous dinosaurs from Mexico. Boletín Geológico y Minero. 126(1), 63-108.
Rivera-Sylva and Longrich, 2024. A new tyrant dinosaur from the Late Campanian of Mexico reveals a tribe of southern tyrannosaurs. Fossil Studies. 2(4), 245-272.

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- 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.
Abler, 1992. The serrated teeth of tyrannosaurid dinosaurs, and biting structures in other animals. Paleobiology. 18(2), 161-183.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen. 210(1), 41-83.
Abler, 2001. A kerf-and-drill model of tyrannosaur tooth serrations. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Life of the Past. Indiana University Press. 84-89.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Life of the Past. Indiana University Press. 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 Osmólska (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.

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 Osmólska (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 für Geologie und Paläontologie 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 Osmólska (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)
Late Cretaceous
Nanxiong Group, Longling Town, Nankang District, 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 Osmólska (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.

Teratophoneini Scherer and Voiculescu-Holvad, 2023 online
Definition- (Teratophoneus curriei <- Albertosaurus sarcophagus, Alioramus remotus, Daspletosaurus torosus, Tyrannosaurus rex) (Scherer and Voiculescu-Holvad, 2023 online)
Reference- Scherer and Voiculescu-Holvad, 2023 online. Re-analysis of a dataset refutes claims of anagenesis within Tyrannosaurus-line tyrannosaurines (Theropoda, Tyrannosauridae). Cretaceous Research. DOI: 10.1016/j.cretres.2023.105780

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 prootic.
(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.

Asiatyrannus Zheng, Jin, Xie and Du, 2024
A. xui Zheng, Jin, Xie and Du, 2024
Etymology- "The generic name is derived from Asia, and the suffix ‘tyrannus’ is derived from the Latin word for 'king' or 'tyrant', to emphasize that this is the new tyrannosaur collected in the continent of Asia. The specific name honors Dr. Xing Xu (Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences), a distinguished dinosaurologist who contributed greatly to the study of dinosaurs from China, including the research of several tyrannosaurs: Guanlong, Dilong, and Yutyrannus. Dr. Xing Xu has also been a great supporter of the paleontological research and science popularization work of the Zhejiang Museum of Natural History."
Late Cretaceous
Nanxiong Group, Shahe Town, Nankang District, Jiangxi, China
Holotype- (ZMNH M30360) (~3.5-4 m; 13+ year old subadult) incomplete skull (475 mm), incomplete mandibles, six partial to incomplete proximal caudal vertebrae, four dorsal proximal chevrons, incomplete right femur (440 mm), incomplete right tibia, left tibial shaft, partial fibulae, partial left metatarsal II, distal right metatarsal III, partial left metatarsal III, right pedal phalanx ?III-1 (~74 mm), distal right metatarsal IV, partial left metatarsal IV, two right pedal phalanges, left (?)metatarsal V fragment
Diagnosis- (after Zheng et al., 2024) small, deep fossa located on lateral surface of premaxilla just lateral to anteroventral border of external naris; large and subrectangular maxillary fenestra; posterior rugosities of nasals form two separated median ridges; accessory jugal horn; ventral margin of anterior jugal process curves ventrally anterior to accessory horn; lineations on ventral postorbital process trend anterodorsally; slender and straight postorbital bar, with almost straight and parallel anterior and posterior margins in lateral view; lateral surangular shelf extends to posterior end.
Comments- This was found in September 2017. Zheng et al. (2024) "interpret the specimen to be a sub-adult that was nearing somatic maturity" based on "a minimum of 13 lines of arrested growth", combined with the fact "The narrower outer zone and the highly developed secondary osteons indicate that the holotype of the Asiatyrannus xui is not a full-growth adult individual, but it already passed through the most rapid growth stage" and "The skull of Asiatyrannus shows several matured cranial morphology."
Using a version of Carr's tyrannosauroid analysis, Zheng et al. (2024) recovered Asiatyrannus as a tyrannosaurine in a trichotomy with Nanuqsaurus and daspletosaurins plus tyrannosaurins.
Reference- Zheng, Jin, Xie and Du, 2024. The first deep‑snouted tyrannosaur from Upper Cretaceous Ganzhou City of southeastern China. Scientific Reports. 14:16276.

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. Papéis 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
= Daspletosaurus "diadematus" Warshaw, Wilson and Fowler, 2022 online
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.
Warshaw, Wilson and Fowler, 2022 online. A transitional species of Daspletosaurus Russell, 1970 from the Judith River Formation of eastern Montana. PeerJ reviewing PDF (2022:07:75847:0:1:NEW)
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 that 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.

Tyrannosaurini Osborn, 1906 sensu Olshevsky, 1995
Definition- (Tyrannosaurus rex <- Albertosaurus sarcophagus, Alioramus remotus, Daspletosaurus torosus, Teratophoneus curriei) (Scherer and Voiculescu-Holvad, 2023 online)
= Tyrannosaurinae sensu Sereno, 1998
Definition- (Tyrannosaurus rex <- Albertosaurus sarcophagus, Daspletosaurus torosus, Gorgosaurus libratus) (modified)
Other definitions- (Tarbosaurus bataar + Tyrannosaurus rex) (Dalman, Loewen, Pyron, Jasinski, Malinzak, Lucas, Fiorillo, Currie and Longrich, 2024)
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. Scherer and Voiculescu-Holvad (2024) and Dalman et al. (2024) give different phylogenetic definitions for this clade, with the former available online in its version of record December 22, 2023 compared to the latter being published online January 11, 2024 (and being in an online-only journal). Of course neither are registered with the ICPN so neither definition is valid under the PhyloCode, and due to ICZN Article 36.1 ("A name established for a taxon at any rank in the family group is deemed to have been simultaneously established for nominal taxa at all other ranks in the family group; all these taxa have the same type genus, and their names are formed from the stem of the name of the type genus [Art. 29.3] with appropriate change of suffix [Art. 34.1]. The name has the same authorship and date at every rank") Tyrannosaurini is credited to Osborn, 1906 anyway so the fact Scherer and Voiculescu-Holvad registered theirs with ZooBank doesn't matter. That being said, Dalman et al.'s definition is far more stable regarding taxonomic content in some recent topologies, as in their topology Bistahieversor and Lythronax would be tyrannosaurins under Scherer and Voiculescu-Holvad's definition, and in Warshaw et al.'s 2024 recent paper on anagenetic Daspletosaurus, D. horneri and D. wilsoni would be tyrannosaurins under that definition. Under Dalman et al.'s definition only Zhuchengtyrannus varies between topologies, and the authorship including several prominent tyrannosaur workers suggests it is more likely to be followed and officially implemented in the future. For now they cover the same taxa in this page's topology, so Scherer and Voiculescu-Holvad's definition is given precedence based on publication timing.
References- Osborn, 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur (Second communication). Bulletin of the American Museum of Natural History. 22(16), 281-296.
Carpenter, 1992. Tyrannosaurids (Dinosauria) of Asia and North America. In Mateer and Chen (eds.). Aspects of nonmarine Cretaceous geology. Ocean Press. 250-268.
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.
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 Osmólska (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.
Dalman, Loewen, Pyron, Jasinski, Malinzak, Lucas, Fiorillo, Currie and Longrich, 2024. A giant tyrannosaur from the Campanian-Maastrichtian of southern North America and the evolution of tyrannosaurid gigantism. Scientific Reports. 13:22124.
Scherer and Voiculescu-Holvad, 2024 (2023 online). Re-analysis of a dataset refutes claims of anagenesis within Tyrannosaurus-line tyrannosaurines (Theropoda, Tyrannosauridae). Cretaceous Research. 155, 105780.

unnamed tyrannosaurin (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 tyrannosaurin 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 tyrannosaurin (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) (Osmólska 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)
Late Maastrichtian, Late Cretaceous
Quipa Formation, Henan, China
?(IVPP V4733; holotype of Tyrannosaurus luanchuanensis) five teeth (A- ?x35.4x?, B- ?x35.1x?, C- ?x36.9x? mm), partial vertebra (Dong, 1979)
Late Campanian-Early Maastrichtian, Late Cretaceous
Dongyuan Formation, Hudieling, Dongyuan County, Heyuan City, Guangdong, China
?(HYMV-7) incomplete tooth (>100x41.79x32.34 mm) (Lü, Liu, Huang, Yuan and Huang, 2009)
Early Maastrichtian?, Late Cretaceous
Zhenshui Formation?, Nanxiong Group, Nanxiong County, Guangdong, China
?(IVPP coll.; unassociated) third premaxillary tooth, lateral tooth (72 mm), dorsal centrum, fragmentary pedal elements (Dong, 1979)
Late Cretaceous
Nanxiong Group, Nankang District, Ganzhou, Jiangxi, China
?(NHMG 8501) lateral tooth (~76x40.4x29 mm) (Mo and Xu, 2015)
Campanian-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)
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. Osmólska 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 Quipa 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 (6-11 per 5 mm mesial, 5.5-10 distal), different from Daspletosaurus and albertosaurines. Since Tarbosaurus has the same density as Tyrannosaurus (Hurum and Sabath, 2003), and the Quipa Formation is Maastrichtian like the Nemegt Formation, it is provisionally considered a junior synonym of T. bataar although its seemingly later age may indicate a distinct species.
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 Formation, suggesting they are not from T. bataar at least.
Dong (1979) referred fragmentary IVPP remains from the Nanxiong Group of Guangdong to Tarbosaurus sp., noting (translated) "Specimens were not collected from the same locality but they are provisionally assigned the same genus based upon morphology." The lateral tooth is said to have 7-8 serrations per 5 mm mesially and 11 per 5 mm distally. When Russell et al. (1993) stated "Rare skeletal remains in the Pingling Formation indicate the presence of ... tyrannosaurids" they cite Dong, 1979 and mean these specimens, as they were using Zhao et al's 1991 stratigraphy where the Pingling Formation is used for the highest section of the Nanxiong Formation (equivalent to the Zhenshui Formation in Li et al.'s 2007 stratigraphy). Their stratigraphic level (as Tarbosaurus sp.) is shown in Figure 2 section 10 number 12 of Mateer and Chen (1992). Note Le Loeuff (2012) places these remains in the Yuanpu Formation (below the Pingling Formation in Zhao et al.'s stratigraphy) and mistakes Russell et al.'s Pingling reference as different tyrannosaurid material. Mo and Xu (2015) described a tooth from the Nanxiong Group of Jiangxi which they referred to Tyrannosauridae indet., noting it was comparable in size to Zhuchengtyrannus but more recurved. It has 7.5-10 serrations per 5 mm distally. Lü et al. (2009) described a tyrannosaurid tooth from the Dongyuan Formation of Guangdong (HYMV-7), where (translated) most of the serrations are missing." All examples are much larger than the contemporaneous Alioramus (Qianzhousaurus holotype) or Asiatyrannus, with NHMG 8501 and HYM-7 at least having a Crown Basal Ratio in the range of Tyrannosaurus and much higher than Alioramus (unreported for Asiatyrannus). Given this and the age of Dong's specimens at least (Early Maastrichtian according to Xi et al. 2021, with the others potentially being that age), they are provisionally retained in Tarbosaurus here.
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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Hurum and Currie, 2000. The crushing bite in tyrannosaurids. Journal of Vertebrate Paleontology. 20(3), 619-621.
Matsumoto, Hashimoto and Sonoda, 2000. Report of preparation works for Mongolian specimens in HMNS from March 1994 to December 1998. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 113-127.
Suzuki and Watabe, 2000a. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1995. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 45-57.
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Carpenter and Smith, 2001. Forelimb osteology and biomechanics of Tyrannosaurus rex. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University Press. 90-116.
Currie, 2001. Nomadic Expeditions, Inc. report on fieldwork in Mongolia, September 2000. Alberta Palaeontological Society, Fifth Annual Symposium. 12-16.
Currie and Dong, 2001. New information on Shanshanosaurus huoyanshanensis, a juvenile tyrannosaurid (Theropoda, Dinosauria) from the Late Cretaceous of China. Canadian Journal of Earth Sciences. 38(12), 1729-1737.
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Hu, Cheng, Pang and Fang, 2001. Shantungosaurus giganteus. Geological Publishing House. 139 pp.
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Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
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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 Zhuchengtyrannus.
References- Gilmore, 1933. Two new dinosaurian reptiles from Mongolia with notes on some fragmentary specimens. American Museum Novitates. 679, 1-20.

undescribed tyrannosaurin (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.

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. mcraeensis Dalman, Loewen, Pyron, Jasinski, Malinzak, Lucas, Fiorillo, Currie and Longrich, 2024
Etymology- "The species name, mcraeensis, refers to the McRae Group of western New Mexico."
Late Campanian-Early Maastrichtian, Late Cretaceous
Staton-LaPoint Tyrannosaurus locality NMMNH 343, Hall Lake Formation of the McRae Group, Sierra County, New Mexico, US
Holotype- (NMMNH P-3698; = NMMNH P-1013-1) maxillary fragments, right postorbital (~411 mm tall), right squamosal, incomplete right palatine, incomplete left dentary (~900 mm), incomplete right splenial, incomplete right angular, incomplete right prearticular, left articular, six incomplete lateral teeth (?x~56x~37, ?x~54x~40, ~89x~39x? mm), incomplete proximal chevrons 939
Diagnosis- (after Dalman et al., 2024; autapomorphies only) dentary very shallow posteriorly and with convex ventral margin (also straight to convex in. T. rex MOR 008); splenial with anteriorly positioned apex (over myohyloid fenestra); splenial with shelf-like dentary overlap ventral to myohyloid fenestra; prearticular weakly bowed.
Comments- Discovered in April 1983, this specimen was originally published by Lozinsky et al. (1984), who referenced "an incomplete jaw of Tyrannosaurus rex" and further stated "This very significant specimen is in the collections of the New Mexico Museum of Natural History and has been made available to us through the courtesy of David Gillette, who first recognized its significance. A detailed description of this specimen will be undertaken by the New Mexico Museum of Natural History and the New Mexico Bureau of Mines and Mineral Resources in a separate paper." Gillette et al. (1986) described it as a Tyrannosaurus rex specimen, although not based on explicit character evidence- "Comparison with the lower jaw of the type specimen of Tyrannosaurus rex (American Museum of Natural History) shows the NMMNH P-1013-1 to be referable to T. rex. This assignment includes the dentary, prearticular, and probably the chevron, which was closely associated with the other elements and closely resembles the anterior chevron of T. rex described by Osborn (1912: pl. 27)." While Lehman and Carpenter (1990) wrote "Other fragmentary specimens, referred to Tyrannosaurus, from elsewhere in New Mexico (Gillette et al., 1986) and Texas (Lawson, 1976) may belong to a new genus (Carpenter, in press b)", the resulting Carpenter 1990 publication only mentioned the Texas specimen TMM 41436-1. Dalman et al. (2024) published a redescription based in part on newly found material (maxillary fragments, postorbital, squamosal, splenial, angular, articular, two additional teeth, additional chevrons), reidentifying the original supposed incomplete articular as a palatine. Note early versions of the pdf incorrectly give the article number as "(2023) 13:22124" despite saying it was published online January 11, 2024. Future versions corrected this to "(2024) 14:22124". Dalman et al. incorrectly state the palatine is from the left side in their materials list, contra the supplementary info and figure S5. While "associated chevrons" are listed in the holotype material, they are not commented on further so it is uncertain how many more were found than the one described by Gillette et al..
Dalman et al. used Loewen's tyrannosauroid analysis to recover Tyrannosaurus mcraeensis sister to T. rex, while it was in a trichotomy with T. rex and Tarbosaurus in Carr's analysis as they had not added characters specific to the mcraeensis + rex clade to that one.
References- Lozinsky, Hunt, Wolberg and Lucas, 1984. Late Cretaceous (Lancian) dinosaurs from the McRae Formation, Sierra County, New Mexico. New Mexico Geology. 6(4), 72-77.
Gillette, Wolberg and Hunt, 1986. Tyrannosaurus rex from the McRae Formation (Lancian, Upper Cretaceous), Elephant Butte Reservoir, Sierra County, New Mexico. New Mexico Geological Society Annual Field Conference Guidebook. 37, 235-238.
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.
Dalman, Loewen, Pyron, Jasinski, Malinzak, Lucas, Fiorillo, Currie and Longrich, 2024. A giant tyrannosaur from the Campanian-Maastrichtian of southern North America and the evolution of tyrannosaurid gigantism. Scientific Reports. 13:22124.
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 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; 13 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; 15 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/)
(Natural History Museum Abu Dhabi coll.; = 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)
(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 (d9 148 mm), 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-Brañas, Torres-Rodríguez, Reyes Luna, González and González-León, 2014)
(ERNO 8550) tooth (41 x 33 x 19 mm) (Serrano-Brañas, Torres-Rodríguez, Reyes Luna, González and González-León, 2014)
(ERNO 8551) tooth (35 x 22 x 17 mm) (Serrano-Brañas, Torres-Rodríguez, Reyes Luna, González and González-León, 2014)
(ERNO 8552) tooth (39 x 27 x 19 mm) (Serrano-Brañas, Torres-Rodríguez, Reyes Luna, González and González-León, 2014)
(ERNO 005) (juvenile?) tooth (28 x 16 x 9 mm) (Serrano-Brañas, Torres-Rodríguez, Reyes Luna, González and González-León, 2014)
(ERNO 006) (juvenile?) tooth (37 x 21 x 12 mm) (Serrano-Brañas, Torres-Rodríguez, Reyes Luna, González and González-León, 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
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
(KUVP 156375) (juvenile) partial premaxilla, maxillae, jugal, postorbital, vomer, ectopterygoid, cervical vertebra, dorsal vertebra, ribs, chevrons, metatarsal I, pedal ungual I, metatarsal IV, two pedal phalanges, distal pedal phalanx, two distal pedal unguals, metatarsal V (Burnham et al., 2018)
(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. Woodward et al. (2020) performed a histological analysis and determined BMRP 2002.4.1's age to be at least thirteen years old. 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 specimen slightly larger than BMRP 2002.4.1 nicknamed Petey (BMRP 2006.4.4) was estimated at over fifteen years old by Woodward et al. (2020). Burnham et al. (2018) report a juvenile Tyrannosaurus from Garfield County, Montana with juvenile bone surface texture that is "nearly the same size as ... BMRP 2002.4.1." Interestingly, "the maxilla ... only contains 12 alveoli" and "the first maxillary tooth of KUVP 156375 is morphologically distinct from that of BMRP 2002.4.1", suggesting this specimen reached skeletal maturity in these features more quickly than the three other known 'Nanotyrannus' specimens with maxillae or supporting the latter being taxonomically distinct from KUVP 156375 and other Tyrannosaurus. 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. As detailed by Greshko (2022), the BHI sold Stan on October 5, 2020 to the Natural History Museum Abu Dhabi, slated to open in 2025.
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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