Tyrannosauroidea Osborn, 1906 sensu Walker, 1964
Definition- (Tyrannosaurus rex <- Passer domesticus) (modified from Sereno, 1998)
Other definitions- (Tyrannosaurus rex <- Ornithomimus velox) (modified from Padian et al., 1999)
(Tyrannosaurus rex <- Allosaurus fragilis, Ornithomimus velox, Deinonychus antirrhopus) (Holtz, 2004)
(Tyrannosaurus rex <- Ornithomimus edmontonicus, Troodon formosus, Velociraptor mongoliensis) (Sereno et al., 2009)
= Deinodontia Flower, 1929
= Deinodontoidea Cope, 1866 emmend. Brown, 1914 sensu Tatarinov, 1964
= Tyrannosauria Olshevsky, 1995
= Tyrannosauroidea sensu Padian et al., 1999
Definition- (Tyrannosaurus rex <- Ornithomimus velox)
= Tyrannosauroidea sensu Holtz, 2004
Definition- (Tyrannosaurus rex <- Allosaurus fragilis, Ornithomimus velox, Deinonychus antirrhopus)
= Tyrannosauridae sensu Holtz, 2004
Definition- (Tyrannosaurus rex <- Eotyrannus lengi)
= Tyrannosauroidea sensu Sereno et al., 2009
Definition- (Tyrannosaurus rex <- Ornithomimus edmontonicus, Troodon formosus, Velociraptor mongoliensis)
Ex-Tyrannosauroidea- Lapparent (1960) suggested Carcharodontosaurus was more closely related to Tyrannosaurus than to megalosaurids/allosaurids, which has been suggested by several other authors since (Paul, 1988; Kurzanov, 1989; Molnar et al., 1990), though it is now recognized as a carnosaur (Sereno et al., 1996). Kurzanov also felt that Diplotomodon was a relative of Carcharodontosaurus, though it has not been restudied recently and is Theropoda indet. on this site. It was however listed as Tyrannosauroidea indet. by Holtz (2004) without comment.
When he named it, Walker (1964) included several taxa in Tyrannosauroidea that are not currently assigned to that clade. Ornithosuchus and Teratosaurus are crurotarsans (Sereno and Arcucci, 1990; Galton, 1985), while Sinosaurus is supposedly a dilophosaurid (Dong, 2003). 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 spinosauroid basal tetanurine (Sereno et al., 1996) and Becklespinax (= Altispinax of Walker) is a carnosaur (Naish, DML 2004; Holtz, 1994). Acrocanthosaurus was often thought to be intermediate between Allosaurus and tyrannosaurids (Walker, 1964; Paul, 1988; Kurzanov, 1989; Bakker et al., 1992), 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 Averianov and Sues (2012)- manual ungual CCMGE 431/12457, femora including CCMGE 724/12457, a tibia, astragali including CCMGE 447/12457 and 448/12457, metatarsals and pedal unguals CCMGE 610/12457 and 609/12457.
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 Tyrannosauridae by Holtz (2004) and Miyashita (2011). Yet Sues and Averianov (2004) and Longrich and Currie (2009) believe it to be a dromaeosaurid. Nessov (1995) listed the Bissekty maxillary fragment N 600/12457 as tyrannosaurid, but Averianov and Sues (2012) reidentify it as dromaeosaurid.
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, that would be defined as tyrannosauroids using the current definition. Besides Acrocanthosaurus and Indosuchus, he also included Chilantaisaurus (a more primitive avetheropod) and Shaochilong (his Chilantaisaurus maortuensis; which is now considered a carcharodontosaurid). 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. It is more probably a ceratosaur (Carrano and Sampson, 2007). Paul also suggested Erectopus was close to tyrannosaurids, but is is more probably a basal tetanurine based on unpublished data. 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, but there has been no evidence presented to show it isn't another kind of tyrannoraptoran. "Tonouchisaurus" was stated to be a tyrannosauroid by the press, but has not been described yet and may be another variety of coelurosaur. Coelurus and Tanycolagreus were both found to be basal tyrannosauroids by Senter (2007), but are here placed slightly closer to birds. Eotyrannus has been assigned to Tyrannosauroidea since it was first announced (e.g. Hutt et al., 2001), but is here placed slightly closer to birds. Dilong and Guanlong have also been assigned to Tyrannosauroidea by most authors (e.g. Xu et al., 2004; 2006), but are here placed slightly closer to birds. Recently, Calamosaurus, Proceratosaurus and Mirischia have been proposed to be tyrannosauroids (Naish, online 2006; Rauhut and Milner, 2008), but their placement depends on the placement of better known taxa like Guanlong and Dilong. Bagaraatan was found to be the basalmost tyrannosauroid by Holtz (2004), but may be more parsimoniously closer to compsognathids. If Coelurus, Tanycolagreus, Eotyrannus, Dilong, Calamosaurus, Proceratosaurus, Mirischia and/or Guanlong are tyrannosauroids, it is likely Nuthetes and Sinocalliopteryx may also belong to that clade. The recent descriptions of Proceratosaurus, Kileskus, Xiongguanlong, Stokesosaurus langhami, Raptorex and Sinotyrannus including new tyrannosauroid characters and codings for relevent taxa may lead to their assignment to Tyrannosauroidea in future updates.
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- 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.
Kurzanov, 1976. Braincase structure in the carnosaur Itemirus n. gen., and some aspects of the cranial anatomy of dinosaurs. Paleontological Journal. 1976, 361-369.
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.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 111-136.
Sues and Averianov, 2004. Dinosaurs from the Upper Cretaceous (Turonian) of Dzharakuduk, Kyzylkum Desert, Uzbekistan. Journal of Vertebrate Paleontology. 24(3), 51A-52A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
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.
Miyashita, 2011. Cranial morphology of the basal tyrannosauroid Itemirus medullaris and evolution of the braincase pneumaticity in non-avian coelurosaurs. Journal of Vertebrate Paleontology. SVP 2011 abstracts, 159A.

Tyrannosauridae sensu Brochu, 2003
Definition- (Alectrosaurus olseni + Gorgosaurus libratus + Albertosaurus sarcophagus + Daspletosaurus torosus + Alioramus remotus + Tarbosaurus bataar + Tyrannosaurus rex)

Alectrosaurus Gilmore, 1933
A. olseni Gilmore, 1933
= Albertosaurus olseni (Gilmore, 1933) Paul, 1988
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Holotype- (AMNH 6554) two 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)
Referred- ?(IGM coll.) several specimens (Currie, 2001)
?(IVPP 170788104) teeth (Currie, Rigby and Sloan, 1990)
?(IVPP 180788-104) teeth (Currie and Eberth, 1993)
Diagnosis- (after Carr, 2005) 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- 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 (2005) found these specimens differ from the holotype and could find no support for their referral.
Currie et al. (1990) report recently discovered Iren Dabasu teeth are identical to Judith River juvenile tyrannosaurid ("Aublysodon") teeth, and that Perle (pers. comm., 1989) referred them to Alectrosaurus. Currie and Eberth (1993) state that Perle (pers. comm. 1989) is studying recently discovered postcranial specimens and report the premaxillary teeth lack serrations (Perle pers. comm. 1989; IVPP 180788-104). Several partial, but undescribed, specimens referred to Alectrosaurus are in the GIN and another was recently collected from Erenhot, China (Currie, 2001). 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.
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, 2005). 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 (2005) 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- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu Formation. Bulletin American Museum of Natural History. 67, 23-78.
Perle, 1977. On the first discovery of Alectrosaurus (Tyrannosauridae, Theropoda)from the Late Cretaceous of Mongolia [in Russian ]. Problemy Geologii Mongolii. 3, 104-113.
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, New York. 107-125.
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.
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.
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.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.

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)
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 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 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.
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, 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. DOI: 10.1002/jmor.10724

"Juratyrant" Brusatte and Benson, in press
"J." langhami (Benson, 2008) Brusatte and Benson, in press
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- Brusatte and Benson (in press) separated langhami from Stokesosaurus, as the synapomorphies were found to be more widely distributed and it claded with Eotyrannus in their tree. Though the online version of the paper appeared in February 2012, it has yet to be published.
References- 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.
Brusatte and Benson, in press. 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

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 [Two dinosaurian vertebrae from the Lower Cretaceous of the Transcapsian steppes]. Zapiski Rossiyskogo Mineralogicheskogo Obshchestva. 60(2, number 1), 110-113.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". in Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 306-317.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Currie, 2000. Theropods from the Cretaceous of Mongolia. In Benton, Shishkin, Unwin and Kurochkin (eds). The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, Cambridge. 434-455.

Xiongguanlong Li, Norell, Gao, Smith and Makovicky, 2010
= "Xiongguanlong" Li, Norell, Gao, Smith and Makovicky, 2009 online
X. baimoensis Li, Norell, Gao, Smith and Makovicky, 2010
= "Xiongguanlong baimoensis" Li, Norell, Gao, Smith and Makovicky, 2009 online
Aptian-Albian, Early Cretaceous
White Ghost Castle field area, 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.
Reference- 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.

Dryptosauridae Marsh, 1890
Dryptosaurus Marsh, 1877
= Laelaps Cope, 1866 (preoccupied Koch, 1836)
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, maxillary tooth, dentary fragment, surangular fragment, two dentary teeth, mid caudal vertebra, mid caudal vertebra (115 mm), mid caudal vertebra (115 mm), distal caudal vertebra (118 mm), distal caudal vertebra (118 mm), distal caudal vertebra (113 mm), distal caudal vertebra (108 mm), distal caudal vertebra (104 mm), distal caudal vertebra (72 mm), two 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)
Referred- ?(AMNH 7624) tooth (Lydekker, 1888)
?(MAPS A1226a) caudal vertebra (Lydekker, 1888)
?(NJSM 14256) tooth (Gallagher, 1993)
? material (Gallagher, Paris and Spamer, 1986)
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 is absent.
(after Brusatte et al., 2011) combination of a reduced humerus (humerus: femur 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 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. A tooth (AMNH 7624) is listed on the AMNH website as coming from the type locality as well.
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.
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.
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 gigantic dinosaur. Am. J. Sci. (ser. 3)14: 87-88.
Osborn, 1898. Paleontological problems. Science. 2, 145-147.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte [The fossil reptile order Saurischia, their development and history]. 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. Simon & Schuster, New York.
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. Unpublished PhD dissertation. 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? macropus (Cope, 1868) Hay, 1902
= Laelaps macropus Cope, 1868
Late Campanian-Early Maastrichtian, Late Cretaceous
Navesink Formation, New Jersey, US
Syntypes- (AMNH 2550) proximal tibia, distal tibia (100 mm wide)
....(AMNH 2551) phalanx II-1 (109 mm), phalanges III-2 (93, 96 mm)
....(AMNH 2552) distal metatarsal IV
....(AMNH 2553) proximal metatarsal II
Diagnosis- (proposed) lateral tibial malleolus at same level as medial malleolus; paired proximoventral processes on pedal phalanges II-1 and III-2.
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. He 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.
The material 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 OMNH 10131), with a laterally square edge as in Appalachiosaurus, Albertosaurus and Tyrannosaurus but unlike Dryptosaurus. The proximal metatarsal is II, and has a sharper posterior corner, more rounded anteromedial corner and shallower lateral notch than Alectrosaurus, Appalachiosaurus and Tyrannosaurus. While ornithomimids also have sharp posterior corners, their proportions are more stout and they lack a lateral notch. The distal metatarsal is IV, and is mostly distinguished from Appalachiosaurus and Tyrannosaurus by its lack of a posterior flange for metatarsal III, and by its somewhat more vertically angled articular surface. These are more similar to the condition in Dryptosaurus and Alectrosaurus. The three pedal phalanges seem to represent II-1 and two III-2's. Phalanx II-1 has a more medially deflected lateral condyle, unlike Gorgosaurus and similar to but more extreme than Alectrosaurus. It has a straighter medial edge than other tyrannosaurs as well. Phalanx II-2 is similar to other tyrannosaurids except in having concave proximal side edges and having paired ventral proximoventral projections. All phalanges are more slender than other tyrannosaurs, though they are more robust than similar-sized ornithomimids (e.g. Gallimimus' holotype). The resemblences to Dryptosaurus seem strongest, which makes sense considering its provenance. It is here considered to be a dryptosaur, though it seems distinct from D. aquilunguis.
References- Leidy, 1865. Memoir of the extinct reptiles of the Cretaceous formations of the United States. Smithsonian Contributions to Knowledge. 14, 1-135.
Cope, 1868. On the genus Laelaps. American Journal of Science. 2(66), 415-417.
Cope, 1870. Synopsis of the extinct Batrachia, Reptilia and Aves of North America. Transactions of the American Philosophical Society. 14, 1-252.
Hay, 1902. Bibliography and Catalogue of the Fossil Vertebrata of North America. Bulletin of the United States Geological Survey. 179, 1-868.
Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History. 46(6), 367-385.
Gallagher, 1997. When Dinosaurs Roamed New Jersey. 176 pp.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds). The Dinosauria (second edition). University of California Press, Berkeley. 111-136.
D? sp. indet. (Baird and Horner, 1979)
Campanian, Late Cretaceous
Black Creek Formation, North Carolina, US
Material- (ANSP 15330) distal femur
(ANSP 15332) two teeth
Comments- Baird and Horner (1979) referred these to Dryptosaurus, though this is not confirmed, as they are also similar to albertosaurines.
Reference- Baird and 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.
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 Camopanian, Late Cretaceous
Basal Marshalltown Formation, New Jersey, US
Material- (NJSM 12436) tooth (Gallagher, 1993)
(NJSM 13087) long 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 14664; cast) tooth
(NJSM 14504) (Gallagher, 1993)
(NJSM 14682) proximal manual phalanx (Gallagher, 1993)
Comments- 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. (Gallagher, 1993)
Early Maastrichtian, Late Cretaceous
Wenonah Formation, New Jersey, US
Material- (MAPS 12106)
Comments- This material was called cf. Dryptosaurus by Gallagher (1993), but may be Appalachiosaurus or another large theropod.
Reference- Gallagher, 1993. The Cretaceous/Tertiary mass extinction event in the northern Atlantic Coastal Plain. The Mosasaur. 5, 75-154.
D? sp. indet. (Baird and Horner, 1979)
Late Cretaceous
Marl Pits of James King, North Carolina, US
Material- (USNM 7189) two femora
Comments- Baird and Horner (1979) referred these to Dryptosaurus, though this is not confirmed, as they are also similar to albertosaurines.
Reference- Baird and 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.
D? sp. indet. (Estes, 1964)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming
Material- teeth
Comments- Originally referred to cf. Dryptosaurus sp. by Estes (1964), based on provenance, these are probably not Dryptosaurus and may be Tyrannosaurus or dromaeosaurid instead.
Reference- Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming. University of California Publications in Geological Sciences. 49, 1-180.

undescribed possible dryptosaur (Sereno, online 2001)
Early Cretaceous
Mazongshan, Inner Mongolia, China
Material- forelimb elements, manual ungual (127 mm)
Comments- This undescribed material was discovered by Sereno's team, during Dinosaur Expedition 2001. 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- 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.
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.
References- 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.
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.

Tyrannosauroidea incertae sedis

Aviatyrannis Rauhut, 2003
A. jurassica Rauhut, 2003
Kimmeridgian, Late Jurassic
Guimarota Formation, Portugal
Holotype- (IPFUB Gui Th 1) ilium (~90 mm)
Referred- (IPFUB Gui Th 2) fragmentary ilium (Rauhut, 2003)
(IPFUB Gui Th 3) proximal ischium (Rauhut, 2003)
?(IPFUB GUI D 89-91) three premaxillary teeth (~6.19 mm) (Zinke, 1998)
?(IPFUB GUI D 174-186) thirteen maxillary and dentary teeth (~10.15 mm) (Zinke, 1998)
Comments- Including Aviatyrannis in Brusatte et al.'s (2010) tyrannosauroid matrix results in a position less derived than Dryptosaurus.
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. 75-82. In Martin and Krebs (eds.). Guimarota - A Jurassic Ecosystem. Verlag Dr. Friedrich Pfeil, Munchen.
Rauhut, 2003. A tyrannosauroid dinosaur from the Upper Jurassic of Portugal. Palaeontology. 46, 903-910.
Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms. Science. 329, 1481-1485.

undescribed possible Tyrannosauroidea (Bakker, 1998)
Tithonian, Late Jurassic
Brushy Basin Member of Morrison Formation, Utah, Wyoming, US
Material- (DNM coll.) (Ford and Chure, 2001)
(WDIS 539) premaxillary tooth (Bakker, 1998)
Comments- Referred to Dromaeosauridae by Bakker. May be Stokesosaurus. Bakker's (1998) isolated Morrison Formation tooth is also said here to be from a tyrannosauroid, perhaps from Aviatyrannis, and differs from the Morrison Formation tyrannosauroid teeth reported by Ford and Chure (2001).
References- Bakker, 1998. Dinosaur mid-life crisis: The Jurassic-Cretaceous transition in Wyoming and Colorado. New Mexico Museum of Natural History and Science Bulletin. 14, 67-77.
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 (Naish, DML 2000)
Late Kimmeridgian-Tithonian, Late Jurassic
Tendaguru Formation, Tanzania
Material- (BMNH coll.) premaxillary tooth (~10 mm)
Description- D-shaped; one side serrated, the other not.
Reference- Naish, DML 2000. http://dml.cmnh.org/2000Apr/msg00440.html

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.

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.

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.

unnamed tyrannosauroid (Zanno and Makovicky, 2011)
Late Aptian, Early Cretaceous
Cloverly Formation, Wyoming, US
Material- (FMNH PR 2750) premaxillary tooth (~9 mm)
Reference- 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 possible Tyrannosauridae (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 (Benson, Barrett, Rich and Vickers-Rich, 2010)
Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P186046) pubes (307 mm)
Comments- This is probably the pubis mentioned by Currie et al. (1996) as NMV P186058, which they referred to Ornithomimosauria. It was later described by Benson et al. (2010) as a tyrannosauroid closer to tyrannosaurids than Guanlong, "Juratyrant" or Raptorex.
References- Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia. Alcheringa. 20(1-2), 73-79.
Benson, Barrett, Rich and Vickers-Rich, 2010. A Southern tyrant reptile. Science. 327, 1613.
Herne, Nair and Salisbury, 2010. Comment on "A Southern tyrant reptile". Science. 329(5995), 1013.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from Southern Australia indicates high polar diversity and climate-driven dinosaur provinciality. PLoS ONE. 7(5), e37122.

undescribed Tyrannosauroidea (Kirkland and Parrish, 1995)
Late Albian, Early Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71399) premaxillary tooth (Fiorillo, 1999)
(juvenile) teeth (Kirkland et al., 1997)
Comments- First noted to be tyrannosaurid by Kirkland and Parrish (1995), these were referred to cf. Alectrosaurus sp. by Kirkland et al. (1997) and Cifelli et al. (1999). It is presumably the indet. aublysodontine listed by Kirkland et al. (1998). However, Alectrosaurus teeth are unknown, and those previously referred to the genus from Mongolia, Kazakhstand and Uzbekistan are currently indeterminate basal and/or juvenile tyrannosauroids.
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.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. New Mexico Museum of Natural History Bulletin. 14, 79-89.
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.

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 possible tyrannosauroid (Weishampel, 1990)
Cenomanian, Late Cretaceous
Potomac Formation, New Jersey, US
Comments- Referred to cf. Albertosaurus sp., but too early to be a tyrannosaurid.
Reference- Weishampel, 1990. Dinosaurian distribution. in Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 63-139.

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 [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 and Sues, 2012. Skeletal remains of Tyrannosauroidea (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan. 34, 284-297.

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)
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.

undescribed possible Tyrannosauridae (Gilmore, 1933)
Cenomanian-Santonian, Late Cretaceous
‘Nantienmen’ beds, Hebei, China
Material- (AMNH 2906) (~3 m) partial dorsal vertebrae, incomplete sacrum, partial caudal vertebrae, proximal radius, fragmentary ilia, fragmentary pubes, fragmentary ischia (Gilmore 1933)
?...(AMNH 6592) (~3 m) tooth, partial cervical vertebra, distal ungual, proximal ulna, phalanges, partial pubes (Gilmore 1933)
Comments- These specimens derive from the same locality and horizon and may be from the same individual. The tooth is serrated distally, suggesting a tyrannosauroid or dromaeosaurid when the age is taken into account. The remains are illustrated on the AMNH website, where they are identified as tyrannosaurid. Perhaps it is a juvenile.
References- Gilmore, 1933. Two new dinosaurian reptiles from Mongolia with notes on some fragmentary specimens. American Museum Novitates 679 1-20.
http://paleo.amnh.org/fossil/show.html?cat_num=FR%202906
http://paleo.amnh.org/fossil/show.html?cat_num=FR%206592

undescribed Tyrannosauroidea (Nessov, 1995)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- (CCMGE 432/12457) incomplete dorsal vertebra (93 mm) (Nessov, 1995)
(CCMGE 433/12457-442/12457, except one) nine teeth (Nessov, 1995)
?(CCMGE 445/12457) pedal phalanx IV-1 (Nessov, 1995)
?(CCMGE 463/12457) pedal ungual (Nessov, 1995)
?(CCMGE 464/12457) pedal ungual (Nessov, 1995)
(CCMGE 477/12475) distal caudal vertebra (Nessov, 1995)
(CCMGE 485/12457) anterior lateral tooth (Nessov, 1995)
(USNM 538123) (juvenile) dorsal neural arch (Averianov and Sues, 2012)
(USNM 538131) partial posterior cervical vertebra (71 mm) (Averianov and Sues, 2012)
(USNM 538132) (adult) anterior dorsal neural arch (Averianov and Sues, 2012)
(USNM 538167) pedal ungual II (Averianov and Sues, 2012)
(USNM 538181) manual ungual II (Averianov and Sues, 2012)
(ZIN PH 2/16) maxillary fragment (Averianov and Sues, 2012)
(ZIN PH 15/16) dentary fragment (Averianov and Sues, 2012)
(ZIN PH 105/16) dorsal neural arch fragment (Averianov and Sues, 2012)
(ZIN PH 106/16) dorsal neural arch fragment (Averianov and Sues, 2012)
(ZIN PH 120/16) mid caudal vertebra (48 mm) (Averianov and Sues, 2012)
(ZIN PH 121/16) astragalar fragment (Averianov and Sues, 2012)
(ZIN PH 124/16) pedal ungual (58 mm) (Averianov and Sues, 2012)
(ZIN PH 507/16) distal caudal vertebra (29.7 mm) (Averianov and Sues, 2012)
(ZIN PH 619/16) manual ungual I (Averianov and Sues, 2012)
(ZIN PH 671/16) anterior cervical centrum (79 mm) (Averianov and Sues, 2012)
(ZIN PH 676/16) incomplete maxilla (261 mm) (Averianov and Sues, 2012)
(ZIN PH 677/16) dentary fragment (Averianov and Sues, 2012)
(ZIN PH 679/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 684/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 693/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 695/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 708/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 733/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 734/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 737/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 755/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 756/16) lateral tooth (Averianov and Sues, 2012)
(ZIN PH 1033/16) premaxillary tooth (Averianov and Sues, 2012)
(ZIN PH 1034/16) premaxillary tooth (Averianov and Sues, 2012)
(ZIN PH 1035/16) premaxillary tooth (Averianov and Sues, 2012)
(ZIN PH 1039/16) premaxillary tooth (Averianov and Sues, 2012)
(ZIN PH 1239/16) (juvenile) posterior mandible (Averianov and Sues, 2012)
(ZIN PH 1476/16) proximal caudal vertebra (80.4 mm) (Averianov and Sues, 2012)
(ZIN PH 2296/16) distal quadrate (Averianov and Sues, 2012)
(ZIN PH 2311/16) anterior dorsal centrum (Averianov and Sues, 2012)
(ZIN PH 2312/16) anterior dorsal centrum (Averianov and Sues, 2012)
(ZIN PH 2330/16) (juvenile) frontal (Averianov and Sues, 2012)
(ZIN PH 2333/16) (juvenile) distal quadrate (Averianov and Sues, 2012)
(ZIN PH 2350/16) (adult) posterior mandible (Averianov and Sues, 2012)
(CCMGE 12457 and ZIN PH 16 coll.) several frontals, two posterior mandibles, fifteen premaxillary teeth (5.3-19.2 mm), fifty-seven lateral teeth (to 65.2 mm), dorsal neural arch fragments, few caudal vertebrae (Nessov, 1995; Averianov and Sues, 2012)
(Paleobiol. Laboratory) (Ford and Chure, 2001)
distal femur, pedal phalanx II-1, distal pedal phalanx II-2 (Carr, 2005)
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.
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.
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 [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.
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.
Ford and Chure, 2001. Ghost lineages and the paleogeographic and temporal distribution of tyrannosaurids. Journal of Vertebrate Paleontology. 21(3), 50A-51A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
Averianov and Sues, 2012. Skeletal remains of Tyrannosauroidea (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan. 34, 284-297.

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 [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.

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.

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 [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, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research. 28, 532-544.

undescribed Tyrannosauroidea (Kirkland, Lucas and Estep, 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Material- partial pes (Eaton pers. comm. 1996 to Kirkland, Lucas and Estep, 1998)
teeth? (Kirkland et al., 1998)
? (juvenile) material (Eaton et al., 1999)
? material (Eaton et al., 1999)
Comments- Kirkland et al. (1998) list both indet. Aublysodontinae and indet. Tyrannosaurinae from this formation. Eaton et al. (1999) listed ?Tyrannosauridae indet. and ?Aublysodon sp. from a Santonian possible Straight Cliffs (or Wahweap?) locality.
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Eaton, Diem, Archibald, Schierup and Munk, 1999. Vertebrate paleontology of the Upper Cretaceous rocks of the Markagunt Plateau, southwestern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 323-333.

unnamed tyrannosauroid (Carpenter, 1982)
Late Coniacian-Early Santonian, Late Cretaceous
Eutaw Formation, Mississippi, US
Material- (MMNS VP103) incomplete pedal phalanx III-1
Comments- While Carpenter (1982) thought this was too broad and dorsoventrally compressed to be ornithomimid (comparing it to tyrannosauroids instead), Baird (1986) stated it was identical to ANSP 15319, which they referred to "Coelosaurus" (as Ornithomimus antiquus). Yet ANSP 15319 may be tyrannosauroid or ornithomimid, and MMNS does resemble Gorgosaurus more than Gallimimus in having a proximally extensive raised dorsal condyle and in being more slender. It is here referred to Tyrannosauroidea.
References- Carpenter, 1982. The oldest Late Cretaceous dinosaurs in North America?. Mississippi Geology. 3(2), 11-17.
Baird, 1986. Upper Cretaceous reptiles from the Severn Formation of Maryland. The Mosasaur. 3, 63-85.

undescribed tyrannosauroid (Lucas et al., 1988)
Santonian, Late Cretaceous
Point Lookout Sandstone, New Mexico, US
Material- (NMMNH P-27482) tooth
Reference- Lucas, Hunt and Pence, 1988. Some Late Cretaceous reptiles from New Mexico. Contributions to Late Cretaceous paleontology and stratigraphy of New Mexico Part III, New Mexico Bureau of Mines & Mineral Resources. 122, 49-60.

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 [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.
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 (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 [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.
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 (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.

unnamed Tyrannosauroidea (Schwimmer et al., 1993)
Late Santonian-Middle Campanian, Late Cretaceous
Blufftown Formation, Alabama, Georgia, US
Material- (CCK-83-3-7) metatarsal shaft fragment
(CCK-84-4-7) partial radius
(CCK-84-4-8) partial ulna
(CCK-85-1-2) metatarsal shaft fragment
(CCK-87-5-1) incomplete metatarsal IV
(CCK-90-1-2) fragmentary pedal(?) phalanx
(CCK-90-5-1) metatarsal shaft fragment
(CCK-90-5-2) metatarsal shaft fragment
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.

unnamed tyrannosauroid (Perle, 1977)
Campanian?, Late Cretaceous
Bayan Shiree 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- 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- Perle, 1977. On the first discovery of Alectrosaurus (Tyrannosauridae, Theropoda) from the Late Cretaceous of Mongolia [in Russian ]. 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. Unpublished PhD dissertation. University of Toronto. 1170 pp.

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.

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 (Carr, 2005)
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 (Carr and Williamson, 2000)
Late Campanian, Late Cretaceous
Fruitland Formation, New Mexico, US
Paratype- (NMMNH P-32824) partial lacrimal (Carr, 2005)
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. Most recently, Brusatte et al. (2010) found it to be sister to Appalachiosaurus+Tyrannosauridae.
References- Lehman and Carpenter, 1990. A partial skeleton of the tyrannosaurid dinosaur Aublysodon from the Upper Cretaceous of New Mexico. Journal of Paleontology. 64, 1026-1032.
Carr and Williamson, 1999. A new tyrannosaurid (Theropoda: Coelurosauria) from the San Juan Basin of New Mexico. Journal of Vertebrate Paleontology. 19(3), 36A.
Williamson and Carr, 1999. A new tyrannosaurid (Dinosauria: Theropoda) partial skeleton from the Upper Cretaceous Kirtland Formation, San Juan Basin, New Mexico. New Mexico Geology, Guidebook 43. 26-29.
Carr and Williamson, 2000. A review of Trannosauridae (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. Unpublished PhD dissertation. 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.
Carr and Williamson, in prep. Phylogeny of the Tyrannosauroidea.

Appalachiosaurus Carr, Williamson and Schwimmer, 2005
A. montogomeriensis Carr, Williamson and Schwimmer, 2005
Middle Campanian, Late Cretaceous
Demopolis Formation, Alabama, US
Material- (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
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.
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. JVP 21(3) 99A.
Williamson and Carr, 2001. Dispersal of pachycephalosaurs and tyrannosauroids between Asia and North America. JVP 21(3) 114A.
Carr and Williamson, 2002. Evolution of basal Tyrannosauroidea from North America. JVP 22(3) 41A.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
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. Unpublished PhD dissertation. 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, in prep. Phylogeny of the Tyrannosauroidea.

Tyrannosauridae Osborn, 1906
Definition- (Gorgosaurus libratus + Albertosaurus sarcophagus + Daspletosaurus torosus + Tarbosaurus bataar + Tyrannosaurus rex) (Holtz, 2004)
Other definitions- (Tyrannosaurus rex <- Alectrosaurus olseni, Aublysodon mirandus, Nanotyrannus lancensis) (modified from Sereno, 1998)
(Aublysodon mirandus + Tyrannosaurus rex) (modified from Holtz, 2001)
(Alectrosaurus olseni + Gorgosaurus libratus + Albertosaurus sarcophagus + Daspletosaurus torosus + Alioramus remotus + Tarbosaurus bataar + Tyrannosaurus rex) (Brochu, 2003)
(Tyrannosaurus rex <- Eotyrannus lengi) (Holtz, 2004)
(Gorgosaurus libratus + Albertosaurus sarcophagus + Tyrannosaurus rex) (Sereno et al., 2009)
(Gorgosaurus libratus + Tyrannosaurus rex) (Sereno et al., 2005 vide Brusatte et al., 2011)
= Deinodontidae Cope, 1866 emmend. Brown, 1914
= Aublysodontidae Nopsca, 1928
= Shanshanosauridae Dong, 1977
= Tyrannosauridae sensu Sereno et al. 2009
Definition- (Gorgosaurus libratus + Albertosaurus sarcophagus + Tyrannosaurus rex)
= Tyrannosauridae sensu Sereno et al., 2005 vide Brusatte et al., 2011
Definition- (Gorgosaurus libratus + Tyrannosaurus rex)
Comments- While unnamed specimens from the Campanian and Maastrichtian are listed below as tyrannosaurids, the presence of taxa such as Alectrosaurus and Dryptosaurus suggests many may be more basal tyrannosauroids.
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 all tyrannosauroids if Eotyrannus is outside that clade as I prefer here. 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.
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 basedon generically unidentifiable material", but this is not a rule in the ICZN.
References- Osborn, 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur (Second communication). Bulletin of the American Museum of Natural History. 22(16), 281-296.
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.

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 in Currie et al., 1990) 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 (Smith, 2005) and is between dentary teeth 4 and 6 of Gorgosaurus in crown compression (Smith, 2002). 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.

Deinodontinae Brown, 1914 sensu Matthew and Brown, 1922
Deinodon Leidy, 1856
D. horridus Leidy, 1856
= Megalosaurus 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)
Late Campanian, Late Cretaceous
Judith River Group, Alberta, Canada
(CMN coll.) several teeth (to 90 mm), metatarsal metatarsal fragments, several phalanges, unguals (Lambe, 1902)
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 to create the short lived combination Megalosaurus 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) noted 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 (e.g. Carr and Williamson, 2004), 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 (Carr and Williamson, 2004).
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. Proc Acad. Nat. Sci. 8(2), 72.
Leidy, 1857.
Leidy, 1860. Extinct vertebrata from the Judith River and Great Lignite Formations of Nebraska. American Philosophical Society Transactions. 11, 139-154.
Cope, 1866. Discovery of a gigantic dinosaur in the Cretaceous of New Jersey Proc. Acad. Nat. Sci. Philadelphia. 18, 275-279.
Cope, 1868.
Leidy, 1868. Remarks on a jaw fragment of Megalosaurus. Proc. Acad. Nat Sci. Philadelphia. 1870, 197-200.
Hay, 1895.
Hay, 1899. On the nomenclature of certain American fossil vertebrates. Am. Geol. 24, 345-349.
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, 1-84.
Matthew and Brown, 1922.
Russell, 1970.
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.
Spamer, Daeschler and Daeschler, 1995. A Study of Fossil Vertebrate Types in the Academy of Natural Sciences of Philadelphia. 434 pp.
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.

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 actual type teeth remain unillustrated.
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.
Osborn, 1902. On Vertebrata of the Mid-Cretaceous of the Northwest Territory. I: Distinctive characters of the Mid-Cretaceous fauna. Contrib. Canad. Pal. III. 1-21.
Olshevsky, 1978. The archosaurian taxa. Mesozoic Meanderings. 1, 1-50.
Glut, 1997. Dinosaurs, the Encyclopedia: Mcfarland & Company, Inc., Publishers, 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- (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.
References- Marsh, 1890. Description of new dinosaurian reptiles. The American Journal of Science. 39, 81-86.
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 [The fossil reptile order Saurischia, their development and history]. Monographien zur Geologie und Palaeontologie. 4(1-2), 1-361.

Deinodon? hazenianus (Cope, 1876) Osborn, 1902
= Laelaps hazenianus Cope, 1876
= Dryptosaurus hazenianus (Cope, 1877) 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(?) (Sankey et al., 2002). The lack of facets or flattened sides eliminates Zapsalis from consideration. Sankey et al. 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. Contrib. Canad. Pal. III. 1-21.
Glut, 1997. Dinosaurs, the Encyclopedia: Mcfarland & Company, Inc., Publishers, 1076 pp.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology. 76(4), 751-763.
Smith, 2005. Heterodonty in Tyrannosaurus rex: Implications for the taxonomic and systematic utility of theropod dentitions. Journal of Vertebrate Paleontology. 25(4), 865-887.

"Ornithomimus" tenuis Marsh, 1890
= Struthiomimus tenuis (Marsh, 1890) Osborn, 1916
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Holotype- (USNM 5814) distal metatarsal III
Comments- Gilmore (1920) illustrated the specimen for the first time and felt it resembled tyrannosaurids more than ornithomimids. Russell (1972) considered Ornithomimus tenuis a possible troodontid, though without comment. However, it is indeed more similar to tyrannosaurids in having an anterior fossa just proximal to the articular condyle and lacking the proximally extended articular surface (posteriorly) of troodontids. It's probably a juvenile Gorgosaurus or Daspletosaurus, based on provenence.
References- Marsh, 1890. Description of new dinosaurian reptiles. The American Journal of Science. Series 3. 39, 81-86.
Osborn, 1916. Skeletal adaptations of Ornitholestes, Struthiomimus, Tyrannosaurus. Bulletin of the American Museum of Natural History. 35(43), 733-771.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. Bulletin of the United States National Museum. 110, 1-154.
Russell, 1972. Ostrich dinosaurs of the Late Cretaceous of Western Canada. Canadian Journal of Earth Sciences. 9, 375-402.

Tarbosaurus? periculosus (Riabinin, 1930) Olshevsky, 1995
= Albertosaurus periculosus Riabinin, 1930
= Deinodon periculosus (Riabinin, 1930) Kuhn, 1965
= Alectrosaurus periculosus (Riabinin, 1930) Olshevsky 1991
= Jenghizkhan periculosus (Riabinin, 1930) Olshevsky, 1995
Coniacian-Maastrichtian, Late Cretaceous
Tsagaan Svita?, Heilongjiang, China
Holotype- (PIN coll.) tooth
References- Riabinin, 1930. [Towards a problem of the fauna and age of the dinosaur beds on the Amur River]. Memoirs of the Russian Mineralogical Society. 59, 41-51.
Kuhn, 1965.
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 [in Japanese]. Kyoryugaku Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.

undescribed Tyrannosauridae (Gilmore, 1933)
Campanian?, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- ?(AMNH 6266) cranial fragments (AMNH online)
(AMNH coll.) pedal elements (Gilmore 1933)
Comments- Gilmore (1933) states that the presence of a tyrannosaurid rivaling Tyrannosaurus in size in the Coniacian-Campanian Iren Dabasu Formation is indicated by a few scattered pedal elements. AMNH 6266 is catalogued as Deinodon? sp., and is thus probably tyrannosaurid.

undescribed Tyrannosauridae (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. JVP 21(3) 50A-51A

undescribed tyrannosaurid (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.
Reference- Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011. A new, large tyrannosaurine theropod from the Upper Cretaceous of China. Cretaceous Research. 32(4), 495-503.

undescribed Tyrannosauridae (Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011)
Campanian, Late Cretaceous
Upper Xingezhuang Formation, Wangshi Series, Shandong, China
Material- (NGMC V287) tooth fragment
(ZCDM coll.) postcrania
Comments- This may belong to Zhuchengotyrannus or the undescribed tyrannosaurid (ZCDM V0030 and V0032), but are not described yet. Note the indeterminate Tyrannosaurus? zhuchengensis is also from the same deposits.
Reference- Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011. A new, large tyrannosaurine theropod from the Upper Cretaceous of China. Cretaceous Research. 32(4), 495-503.

undescribed tyrannosaurid (Ford and Chure, 2001)
Campanian, Late Cretaceous
Maortu, Mongolia
Material- (IVPP V239)
Reference- Ford and Chure, 2001. Ghost lineages and the paleogeographic and temporal distribution of tyrannosaurids. JVP 21(3) 50A-51A

unnamed Tyrannosauridae (Riabinin, 1930)
Campanian-Maastrichtian, Late Cretaceous
Tsagaan Svita, Russia
Material- (1/789) lateral tooth (?x23.6x13.8 mm) (Bolotsky, 2011)
(1/790) lateral tooth (71x?x14.4 mm) (Bolotsky, 2011)
(1/791) lateral tooth (64.9x?x13 mm) (Bolotsky, 2011)
(1/797) premaxillary tooth (?x?x11 mm) (Bolotsky, 2011)
(1/799) lateral tooth (?x14.3x10 mm) (Bolotsky, 2011)
(1/800) lateral tooth (27.5x11.4x11.3 mm) (Bolotsky, 2011)
(1/802) lateral tooth (42x16.3x9.7 mm) (Bolotsky, 2011)
(1/819) lateral tooth (33x16.8x11.6 mm) (Bolotsky, 2011)
(1/820) lateral tooth (33x17.7x10.5 mm) (Bolotsky, 2011)
(1/823) lateral tooth (38x17.9x11.5 mm) (Bolotsky, 2011)
(1/824) lateral tooth (38.6x17.3x9.9 mm) (Bolotsky, 2011)
(1/825) lateral tooth (?x17.2x9.9 mm) (Bolotsky, 2011)
(1/826) lateral tooth (32x17x9.1 mm) (Bolotsky, 2011)
(1/841) lateral tooth (73x26.7x17.5 mm) (Bolotsky, 2011)
(1/842) lateral tooth (59x26.4x21 mm) (Bolotsky, 2011)
(1/846) lateral tooth (30x13.5x8.2 mm) (Bolotsky, 2011)
(1/847) lateral tooth (35.5x15.5x? mm) (Bolotsky, 2011)
(1/848) lateral tooth (36x?x? mm) (Bolotsky, 2011)
(1/849) lateral tooth (29.5x13.6x9.5 mm) (Bolotsky, 2011)
(1/850) lateral tooth (29.3x16.3x10.9 mm) (Bolotsky, 2011)
(1/851) lateral tooth (27.5x16.4x9 mm) (Bolotsky, 2011)
(1/852) lateral tooth (26.5x14.7x8 mm) (Bolotsky, 2011)
(1/1077) lateral tooth (32x15x8.5 mm) (Bolotsky, 2011)
(1/1078) lateral tooth (27x?x8.4 mm) (Bolotsky, 2011)
References- Riabinin, 1930. [On the age and fauna of the dinosaur beds on the Amur River] (in Russian). Mémoir, Société Mineral Russia. 59, 41-51.
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.

unnamed Tyrannosauridae (Bolotsky, 2011)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- (2/10) lateral tooth (33.5x18.6x13.7 mm)
(2/11) lateral tooth (37x19.3x12.2 mm)
(2/13) lateral tooth (?x17.5x12.6 mm)
(2/421) lateral tooth (72x27.9x16.5 mm)
(2/424) lateral tooth (?x19.2x12 mm)
(2/427) lateral tooth (16.5x9.8x6.7 mm)
(2/428) premaxillary tooth (26 mm)
(2/431) lateral tooth (16.5x9.2x6.2 mm)
(2/434) lateral tooth (?x20.7x12.5 mm)
(2/435) lateral tooth (36.5x12.9x11.5 mm)
(2/436) lateral tooth (?x15.4x10.3 mm)
(2/1027) lateral tooth (?x11.4x9.8 mm)
(2/1028) lateral tooth (24x14.5x7.3 mm)
(2/1037) lateral tooth (73x26.5x18 mm)
(2/1038) lateral tooth (46x17.9x11.2 mm)
Reference- 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.

undescribed tyrannosaurid (Russell, Russell and Sweet 1993)
Late Maastrichtian, Late Cretaceous
Pingling Formation, Guandong, China
Reference- Russell, D.A., D.E. Russell, and A.R. Sweet. 1993. The end of the dinosaurian era in the Nanxiong Basin. VPA 31(2): 139-145.

undescribed Tyrannosauridae (Erickson, 1995)
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)
material (Nelms, 1992)
tooth tip (Clos, 2004)
Comments- Most of these are from the Campanian-Maastrichtian Kogosukruk Tongue portion of the Prince Creek Formation, five of which are Early Maastrichtian in age.
May all be the same material. Clos (2004) referred a partial tooth from the Early Maastrichtian to Albertosaurus. This is possible given its age.
References- Erickson, 1992.

undescribed Tyrannosauridae
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
Material- (GSC 8724; = CMN 8724) tooth (Ford and Chure, 2001)
(MR-4:74) tooth (Baszio, 1997)
(RTMP 20021) (juvenile) tooth (Ryan and Russell, 2001)

undescribed tyrannosaurid (Langston, 1960)
Early Campanian, Late Cretaceous
Mooreville Chalk Member of Selma Formation, Alabama, US
Material- (FMNH P27398) pedal phalanx

undescribed Tyrannosauridae (Kirkland, Lucas and Estep, 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah, US
Material- (OMNH 23635) tooth (Parrish, 1999)
(OMNH 24309; in part) (juvenile) tooth (Parrish, 1999)
Comments- Parrish (1999) listed OMNH 24635 as Tyrannosauridae and 24309 as cf. Aublysodon.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

undescribed tyrannosaurid (Parker and Rowley, 1989)
Early Campanian, Late Cretaceous
Blackhawk Formation, Utah
Material- skull, tooth

undescribed Tyrannosauridae
Early Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- teeth (Mongelli and Varricchio, 1998)

undescribed Tyrannosauridae
Early Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- (MOR 414) five teeth (MOR online)
(MOR 1116) tibia, metatarsal (MOR online)

undescribed Tyrannosauridae
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
Material- (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)

undescribed tyrannosaurid
Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
Material- (YPM 9834) (YPM online)

unnamed Tyrannosauridae (Miller, 1967)
Campanian, Late Cretaceous
Black Creek Formation, North Carolina, US
Material- (ANSP 15303) anterior maxilla
(ANSP 15319) pedal phalanx III-3 (62 mm)
(ANSP 15331) tooth
(USNM 7199) tooth
Comments- 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 actually 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.
References- Miller, 1967. Cretaceous vertebrates from Phoebus Landing, North Carolina. Proc. Acad. Nat. Sci. Phila. 779(5), 219-235.
Baird and Horner, 1979. Cretaceous dinosaurs of North Carolina. Brimleyana. 2, 1-28.

undescribed Tyrannosauridae (Lucas et al., 1990)
Campanian, Late Cretaceous
Ringbone Formation, New Mexico, US
Material- (NMMNH P-3050) proximal caudal centrum (Lucas et al., 1990)
(NMMNH P-12997) tooth (Lucas et al., 1990)

undescribed Tyrannosauridae (Westgate, Brown and Pittman, 2002)
Campanian, Late Cretaceous
San Carlos Formation, Mexico
Reference- Westgate, Brown and Pittman, 2002. Discovery of dinosaur remains in coastal deposits near Ojinaga, Mexico. JVP 22(3) 118A-119A.

unnamed Tyrannosauridae (Lehman, 1985)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Material- (LSUMG 489:5580) tooth fragment (Sankey, 2001)
(TMM 42534) (Lehman, 1989)
teeth (Lehman, 1985)
Late Campanian, Late Cretaceous
Aguja Formation Mexico
Material- teeth, limb elements (Westgate et al., 2002)
Comments- 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).
References- Lehman, 1985. Stratigraphy, sedimentology, and paleontology of Upper Cretaceous (Campanian-Maastrichtian) sedimentary rocks in Trans-Pecos, Texas. Unpublished Ph.D. dissertation, University of Texas at Austin, Austin. 299 pp.
Standhardt, 1986. Vertebrate paleontology of the Cretaceous/Tertiary transition of Big Bend National Park, Texas. Unpublished Ph.D. dissertation, Louisiana State University, Baton Rouge. 298 pp.
Lehman, 1989. Chasmosaurus mariscalensis, sp. nov., a new ceratopsian dinosaur from Texas. Journal of Vertebrate Paleontology, 9:137–162.
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, 472-493.
Sankey, 1998. Vertebrate paleontology and magnetostratigraphy of the upper Aguja Formation (Late Campanian), Talley Mountain area, Big Bend National Park, Texas. Unpublished Ph.D. dissertation, Louisiana State University, Baton Rouge. 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.

undescribed Tyrannosauridae (Ford and Chure, 2002)
Late Campanian, Late Cretaceous
El Gallo Formation, Mexico
Material- (IGM 4302; = LACM 20886) (juvenile) tooth
(IGM coll.; = LACM 7253/28999) (juvenile) tooth
(IGM coll.; = LACM 3294/24580) (juvenile) tooth
teeth (Rodriguez, 1999; Rodriguez de la Rosa and Aranda-Manteca, 2000)
material (Hernandez-Rivera, 1997)
Description- These 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 (Rodriguez, 1999).
Comments- Supposedly a potentially new taxon.
References- Hernandez-Rivera, 1997.
Rodriguez, 1999. Theropod teeth from the Late Cretaceous El Gallo Formation, Baja California, Mexico. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
Ford, T. L., and Chure, D. J., 2002, "Aublysodon" teeth from the El Gallo Formation (Late Campanian) of Baja California: the southernmost record of tyrannosauroid theropods: In: Western Association of Vertebrate Paleontologists with Mesa Southwest Museum and Southwest Paleontological Society of Mesa, Arizona, First Meeting of the New Millennium, Mesa Southwest Museum Bulletin n. 8, p. 75-89

undescribed tyrannosaurid (Molnar, 1974)
Campanian, Late Cretaceous
La Bocana Roja Formation, Mexico
Material- (LACM 28237) several teeth, metatarsal

undescribed Tyrannosaurid
Campanian, Late Cretaceous
Trent River Formation, British Columbia, Canada
Material- (lost) vertebra

undescribed Tyrannosauridae
Late Campanian, Late Cretaceous
Foremost Formation, Alberta, Canada
(RTMP 88.86.4) tooth (Ryan and Russell, 2001)
(RTMP 92.30.219) tooth
(RTMP coll.) (juvenile) tooth (Ryan and Russell, 2001)

undescribed Tyrannosauridae (pr= Gorgosaurus libratus or Daspletosaurus sp. nov.)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Material- (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, 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, 1990)
(RTMP 80.8.192) (juvenile) tooth (Molnar and Carpenter 1989)
(RTMP 80.16.485) (juvenile) frontal, tooth (Molnar and Carpenter 1989)
(RTMP 80.16.864) tooth (80 mm) (Currie, 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, 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)

undescribed Tyrannosauridae
Late Campanian, Late Cretaceous
Oldman Formation, Alberta, Canada
Material- (RTMP 96.62.48) (juvenile) tooth (Ryan and Russell, 2001)
(YPM-PU 24515) (YPM online)

undescribed Tyrannosauridae
Late Campanian, Late Cretaceous
Judith River Group, Saskatchewan, Canada
Material- teeth, caudal vertebra (Tokaryk, 1986)

undescribed Tyrannosauridae
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
Material- (AMNH 2479) (Cope, 1876)
(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, 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 (MOR online)
(MOR 769) partial skeleton (MOR online)
(MOR 1029) tooth (MOR online)
(MOR 1061) fragmentary nasal (MOR 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)
material (Fiorillo, 1989)

undescribed Tyrannosauridae
Late Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
Material- (MOR 468) cranial fragment, vertebrae, pelvic element, femur, phalanx (MOR online)
(MOR 553E-6-19-91-69) radius (MOR online)
(MOR 565) tooth (MOR online)
(MOR 589) braincase (MOR online)
(MOR 1130) skeleton (MOR online)

undescribed Tyrannosauridae (Carr and Williamson, 2000)
Late Campanian, Late Cretaceous
Fruitland or Lower Kirtland Formation, New Mexico, US
Material- (LACM 45985) tooth fragments (Carr and Williamson, 2000)
(NMMNH P-22693) partial pedal phalanx (Carr and Williamson, 2000)
(NMMNH P-22908) distal metatarsal II, distal metatarsal III (Carr and Williamson, 2000)
(NMMNH P-27744) distal metatarsal IV, fragments (Carr and Williamson, 2000)
(NMMNH P-27773) digit I (Carr and Williamson, 2000)
(NMMNH P-27787) proximal rib (Carr and Williamson, 2000)
(NMMNH coll.) 19+ teeth (see Carr and Williamson, 2000 for numbers)
(USNM 365551) incomplete pubis, femur (665 mm), tibia, metatarsal (Carr and Williamson, 2000)
Reference- Carr and Williamson, 2000. A review of Trannosauridae (Dinosauria: Coelurosauria) from New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science. Bulletin 17. 113-146.

undescribed Tyrannosauridae (Armstrong-Ziegler, 1980)
Late Campanian, Late Cretaceous
Fruitland Formation, New Mexico, US
Material- (KUVP 85370) limb element (Carr and Williamson, 2000)
(MNA Pl.1623) three teeth (Armstrong-Ziegler, 1980)
(NMMNH P-30077) tooth (Carr and Williamson, 2000)
(NMMNH P-32590) tooth (Carr and Williamson, 2000)
(PMA P 73.30.1) metatarsus (480 mm) (Holtz 1994)
References- Carr and Williamson, 2000. A review of Trannosauridae (Dinosauria: Coelurosauria) from New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science. Bulletin 17. 113-146.

undescribed Tyrannosauridae (Lehman and Carpenter, 1990)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-22976; = UNM B-828?) femur (995 mm) (Lehman and Carpenter, 1990)
(NMMNH P-25073) proximal scapula (Carr and Williamson, 2000)
(NMMNH P-27281) pedal phalanx (Carr and Williamson, 2000)
(NMMNH P-27620) pedal phalanges (Carr and Williamson, 2000)
(NMMNH P-29164) tibia (Carr and Williamson, 2000)
(NMMNH P-30072) partial pedal phalanx (Carr and Williamson, 2000)
(NMMNH P-30074) caudal vertebra (Carr and Williamson, 2000)
(NMMNH P-30075) pedal phalanx fragment (Carr and Williamson, 2000)
(NMMNH coll.) twelve teeth (see Carr and Williamson, 2000 for numbers)
References- Carr and Williamson, 2000. A review of Trannosauridae (Dinosauria: Coelurosauria) from New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science. Bulletin 17. 113-146.

undescribed Tyrannosauridae (Gilmore, 1916)
Late Campanian, Late Cretaceous
De-ne-zin Member of Kirtland Formation, New Mexico, US
Material- (NMMNH 12999, others; =UNM FKK-077, 078, 079, 080) four teeth (Lucas et al., 1987)
(NMMNH P-20879) pedal phalanx (Hunt and Lucas, 1992)
(NMMNH P-25071) distal femur (Carr and Williamson, 2000)
(NMMNH P-25085) tibia (993 mm) (Carr and Williamson, 2000)
(NMMNH P-26276) manual ungual (Carr and Williamson, 2000)
(NMMNH P-27276) distal metatarsal II (Carr and Williamson, 2000)
(NMMNH P-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 coll.) >64 teeth (see Carr and Williamson, 2000 for numbers)
(USNM 8346) dentary (Gilmore, 1916, 1920, 1935)
(USNM 8355) (juvenile) premaxillary tooth (Gilmore, 1916, 1920)
References- Carr and Williamson, 2000. A review of Trannosauridae (Dinosauria: Coelurosauria) from New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science. Bulletin 17. 113-146.

undescribed Tyrannosauridae (Carr and Williamson, 2000)
Late Campanian, Late Cretaceous
Lower Kirtland Formation, New Mexico, US
Material- (KUVP 12164) teeth (Carr and Williamson, 2000)
(NMMNH P-7178) partial phalanx (Carr and Williamson, 2000)
(NMMNH coll.) eight teeth (see Carr and Williamson, 2000 for numbers)
References- Carr and Williamson, 2000. A review of Trannosauridae (Dinosauria: Coelurosauria) from New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science. Bulletin 17. 113-146.

undescribed Tyrannosauridae (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)
(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)
Reference- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

undescribed tyrannosaurid (Lehman, 1985)
Late Campanian, Late Cretaceous
San Carolos Formation, Texas, US

undescribed tyrannosaurid (Young, 1987)
Late Campanian, Late Cretaceous
Williams Fork Formation of Mesaverde Group, Colorado, US
Material- tooth

undescribed tyrannosaurid (Breithaupt, 1985)
Late Campanian, Late Cretaceous
Mesaverde Group, Wyoming, US
Comments- Originally referred to Albertosaurus, it is likely too early to be that genus.
Reference- Breithaupt, 1985. Nonmammalian vertebrate faunas from the Late Cretaceous of Wyoming. In Thirty-Sixth Annual Field Conference Guidebook (C. F. Nelson, Ed.), pp. 159-175. Wyoming Geological Association, Casper.

undescribed Tyrannosauridae (Konecy, 1994)
Fort Crittenden Formation, Arizona, US
Late Campanian, Late Cretaceous
Material- (Konecy private coll.) two tooth fragments (Konecy, 1994)
(UALP 1925; = A 25) tooth (McCord, 1997)
(UALP 1927; = A 26) tooth (McCord, 1997)
(UALP 1928; = A 27) tooth (McCord, 1997)
references- Krzyzanowski, Lucas and Heckert, 2001. Late Campanian (Judithian) vertebrate fauna of the Fort Crittenden Formation, Southeastern Arizona. JVP 21(3) 69A-70A.

undescribed tyrannosaurid (Murry, Boyd, Wolleben and Wilson, 1960)
Late Campanian-Early Maastrichtian (Brinkman et al., 2002), Late Cretaceous
Cerro del Pueblo Formation, Mexico

undescribed tyrannosaurid (Sullivan and Williamson, 1997)
Late Campanian-Maastrichtian, Late Cretaceous
Kirtland Formation, New Mexico, US
Material- (KU 12419) (juvenile) tooth (Molnar and Carpenter, 1989)
(PMU.R40; =R1240?) pedal ungual (Sullivan and Williamson, 1997)

undescribed Tyrannosauridae (Brown, 1910)
Late Campanian-Maastrichtian, Late Cretaceous
Fruitland or Kirtland Formation, New Mexico, US
Material- (AMNH 2479 in part; holotype of Dysganus encaustus) tooth (Carr and Williamson, 2000)
(AMNH coll.) teeth (Brown, 1914)
(KUVP 14958) mandibular fragment (Carr and Williamson, 2000)
(KUVP 15135) tooth (Carr and Williamson, 2000)
(KUVP 15145) teeth (Carr and Williamson, 2000)
(KUVP 15234-15235) two unguals (Carr and Williamson, 2000)
(KUVP 15262) tooth (Carr and Williamson, 2000)
(KUVP 15605) teeth (Carr and Williamson, 2000)
(KUVP 16042) tooth, phalanx (Carr and Williamson, 2000)
(KUVP 96846) femur (Carr and Williamson, 2000)
(KUVP 96861) astragalus (Carr and Williamson, 2000)
(KUVP 96878) tooth (Carr and Williamson, 2000)
(KUVP 96888) mandible (Carr and Williamson, 2000)
(PMU.R35; = R1235?) anterior dentary (Sullivan and Williamson, 1997)
(PMU.R36; = R56?) tooth (Sullivan and Williamson, 1997)
(PMU.R85) partial dentary (Carr and Williamson, 2000)
(USNM 10754) metatarsus (540 mm) (Holtz 1994)
(USNM coll.) teeth (Gilmore, 1916)
(uncollected) (~8 m) vertebrae, limb elements (Brown, 1910)
References- Carr and Williamson, 2000. A review of Trannosauridae (Dinosauria: Coelurosauria) from New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science. Bulletin 17. 113-146.

undescribed Tyrannosauridae (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

undescribed tyrannosaurid (Schneiderman pers. comm. to Ford and Chure, 2001)
Early Maastrichtian?, Late Cretaceous
Holmesville, Nebraska, US
Material- (Nebraska State Museum coll.) partial tooth

undescribed tyrannosaurid (Hoganson, Erickson and Getman, 1994)
Maastrichtian, Late Cretaceous
Timber Lake Member of Fox Hills Formation, North Dakota, US
Material- tooth

undescribed Tyrannosauridae
Maastrichtian, Late Cretaceous
St. Mary River Formation, Alberta, Canada
Material- (CMN 9589) tooth (Langston, 1975)
(CMN 9723) tooth (Langston, 1975)
(CMN 10650) tooth (Langston, 1975)
(CMN 10651) tooth (Langston, 1975)
(CMN 10652) tooth (Langston, 1975)
(CMN 10675) tooth (Langston, 1975)

undescribed Tyrannosauridae
Maastrichtian, Late Cretaceous
Lower Ravenscrag Formation, Saskatchewan, Canada
Material- teeth (Russell, 1930)

undescribed tyrannosaurid
Late Maastrichtian, Late Cretaceous
Denver Formation, Colorado, US
(UCMP 38060) (juvenile) tooth (Molnar and Carpenter, 1989)

undescribed tyrannosaurid
Late Maastrichtian, Late Cretaceous
Livingston Formation, Montana, US
undescribed Tyrannosauridae
Material- (MOR 002) postcranial skeleton (MOR online)

undescribed Tyrannosauridae
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (MOR 064) tooth fragment
(MOR 072) tooth fragment
(MOR 336) centrum (MOR online)
(RTMP 87.112.33) (juvenile) tooth (Molnar and Carpenter 1989)
(RTMP 87.114.7) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124367) (juvenile) tooth (6.1 mm) (Carpenter 1982)
(UCMP 88125, 109015, 109018, 119676, 119677, 120135, 120262, 120306, 120352, 123373, 123508, 123509, 123545, 124486, 12549) teeth (Ford and Chure, 2001)
(UCMP 112003) distal tibia (Ford and Chure, 2001)
(UCMP 119508) phalanges (Ford and Chure, 2001)
(UCMP 119578, 119678, 120080, 120137) tooth fragments (Ford and Chure, 2001)
(UCMP 119579, 119580, 120017, 120048) phalanges (Ford and Chure, 2001)
(UCMP 119725) metatarsal? (Ford and Chure, 2001)
(UCMP 119785) six vertebrae (Ford and Chure, 2001)
(UCMP 119786) two teeth (Ford and Chure, 2001)
(UCMP 119787) two teeth (Ford and Chure, 2001)
(UCMP 119788) three teeth (Ford and Chure, 2001)
(UCMP 119853) (juvenile) four teeth (Ford and Chure, 2001)
(UCMP 119854) two tooth fragments (Ford and Chure, 2001)
(UCMP 119929) seven teeth (Ford and Chure, 2001)
(UCMP 119931) four teeth (Ford and Chure, 2001)
(UCMP 119932) tooth (Ford and Chure, 2001)
(UCMP 119933) tooth (Ford and Chure, 2001)
(UCMP 119934) six tooth fragments (Ford and Chure, 2001)
(UCMP 119935) three caudals. (Ford and Chure, 2001)
(UCMP 119936) five phalanges (Ford and Chure, 2001)
(UCMP 120001) two unguals (Ford and Chure, 2001)
(UCMP 120081) ungual (Ford and Chure, 2001)
(UCMP 120136) two teeth (Ford and Chure, 2001)
(UCMP 120194) three tooth fragments (Ford and Chure, 2001)
(UCMP 120260) two teeth (Ford and Chure, 2001)
(UCMP 120261) six tooth fragments (Ford and Chure, 2001)
(UCMP 120263) caudal vertebra (Ford and Chure, 2001)
(UCMP 120305) two teeth (Ford and Chure, 2001)
(UCMP 120307) pedal phalanx. (Ford and Chure, 2001)
(UCMP 120339) two teeth (Ford and Chure, 2001)
(UCMP 120340) ungual (Ford and Chure, 2001)
(UCMP 120341) caudal vertebra (Ford and Chure, 2001)
(UCMP 120843) two teeth (Ford and Chure, 2001)
(UCMP 120844) two manual phalanges (Ford and Chure, 2001)
(UCMP 120845) four ungual fragments (Ford and Chure, 2001)
(UCMP 123342) eight teeth (Ford and Chure, 2001)
(UCMP 123343) 20+ teeth (Ford and Chure, 2001)
(UCMP 123344) seven teeth (Ford and Chure, 2001)
(UCMP 123345) 20+ teeth (Ford and Chure, 2001)
(UCMP 123346) 40+ teeth (Ford and Chure, 2001)
(UCMP 123347) metatarsal (Ford and Chure, 2001)
(UCMP 123567) three teeth (Ford and Chure, 2001)
(UCMP 124484, cast) tooth (Ford and Chure, 2001)
(UCMP 124485) four teeth (Ford and Chure, 2001)
(YPM 54459) (YPM online)
(YPM 54461) (YPM online)
(YPM 55508) (YPM online)
(YPM 55519) (YPM online)
(YPM 55532) (YPM online)
(YPM 55540) (YPM online)
(YPM 55541) (YPM online)
(YPM 55559) (YPM online)
(YPM 55569) (YPM online)
(YPM 55597) (YPM online)
(YPM 55618) (YPM online)
teeth, bones (Triebold, 1997)
Comments- These are probably Tyrannosaurus, based on their provenance.

undescribed Tyrannosauridae
Late Maastrichtian, Late Cretaceous
Lance Formation, Montana, Wyoming, US
Material- (CMN 12102) femur (McIntosh, 1981)
(CMN 30749) thirteen teeth (McIntosh, 1981)
(UCMP 37878) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 38804) tooth (Ford and Chure, 2001)
(UCMP 39505) tooth (Ford and Chure, 2001)
(UCMP 43447) (juvenile) tooth (Carpenter 1982)
(UCMP 45063) tooth (Ford and Chure, 2001)
(UCMP 73091) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 73101) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 85141) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124237) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124399) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124406) (juvenile) tooth (Carpenter 1982)
(UCMP 124978) (juvenile) tooth (Carpenter 1982)
(UCMP 124979) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124980) (juvenile) tooth (Carpenter 1982)
(UCMP 124981) (juvenile) tooth (7 mm) (Molnar and Carpenter 1989)
(UCMP 124982) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124993) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124994) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124995) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 124996) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 125229) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 125230) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 125233) (juvenile) tooth (Molnar and Carpenter 1989)
(UCMP 125234) (juvenile) tooth (Molnar and Carpenter 1989)
(UW 15219) tooth (Breithaupt, 1982)
(YPM-PU 18156) (YPM online)
Comments- These are probably Tyrannosaurus, based on their provenance.

undescribed Tyrannosauridae (Cope, 1885)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Kirtland Formation, New Mexico, US
Material- (AMNH 2359) tooth, tooth fragments (Cope, 1885)
(AMNH 5882) pedal phalanx (Carr and Williamson, 2000)
(NMMNH P-28367) tooth fragment (Carr and Williamson, 2000)
(NMMNH P-28368) tooth (Carr and Williamson, 2000)
(NMMNH P-28369) tooth fragment (Carr and Williamson, 2000)
(NMMNH P-30076) tooth fragment (Carr and Williamson, 2000)
(NMMNH P-32566) tooth fragments (Carr and Williamson, 2000)
(NMMNH P-32567) tooth (Carr and Williamson, 2000)
(NMMNH P-32588) partial premaxillary tooth (Carr and Williamson, 2000)
(NMMNH P-32598) partial premaxillary tooth (Carr and Williamson, 2000)
(SMP VP coll.) femur (Carr and Williamson, 2000)
(privately owned) metatarsal IV (Lehman, 1981)
References- Carr and Williamson, 2000. A review of Trannosauridae (Dinosauria: Coelurosauria) from New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science. Bulletin 17. 113-146.

undescribed tyrannosaurid (Gilmore, 1919)
Late Maastrichtian, Late Cretaceous
Ojo Alamo Formation, New Mexico, US
Material- (USNM coll.) (~12 m) teeth, vertebrae (Gilmore, 1919)

undescribed Tyrannosauridae (Lehman, 1982)
Late Maastrichtian, Late Cretaceous
Tornillo Group, Texas, US
Material- (TAMU 1372) tooth
(TMM 31201-1) tooth
(TMM 31201-6) tooth
(TMM 31221-1) femur
(TMM 40573-1) tibia
(TMM 41395-3) tooth
(TMM 41541-2) tooth
(TMM 41541-3) tooth
(TMM 42291-1) tooth
(TMM 42291-2) tooth
(TMM 49710-1) tooth
(TMM AM 144) tooth

undescribed Tyrannosauridae (Lehman, 1985)
Late Maastrichtian, Late Cretaceous
Javelina Formation, Texas, US

undescribed Tyrannosauridae (Lehman, 1985)
Late Maastrichtian, Late Cretaceous
El Picacho Formation, Texas, US

undescribed Tyrannosauridae (Gilmore, 1946)
Late Maastrichtian, Late Cretaceous
North Horn Formation, Utah, US
Material- teeth, manual ungual, distal metatarsal II or IV
Comments- These were identified as deinodontids, representing both large and small individuals. It's possible some belong to Tyrannosaurus, which is known from the formation, or other coelurosaurs.
Reference- Gilmore, 1946. Reptilian fauna of the North Horn Formation of central Utah: United States Geological Survey Professional Paper. 210-C, 29-53.

undescribed Tyrannosauridae (Wroblewski, 1998)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
Material- (juvenile) teeth? (Wroblewski, 1998)

Albertosaurinae Currie et al., 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 [in Japanese]. Kyoryugaku Saizensen (Dino Frontline). 9, 92-119; 10, 75-99.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. 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, Saskatchewan, Canada; Judith River Group, Montana, US
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 3963) dentary
(AMNH 5218) femur (954 mm), tibia (850 mm), metatarsus (515 mm)
(AMNH 5232) metatarsus (560 mm)
(AMNH 5233) metatarsus (480 mm)
(AMNH 5234) metatarsus (440 mm)
(AMNH 5235) femur (870 mm), metatarsus (510 mm)
(AMNH 5346) maxilla (Carr, 1996)
(AMNH 5423) anterior maxilla, skull roof, dentary, skeleton including femur (600 mm), tibia (630 mm), metatarsus (440 mm)
(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), phalanges (Carr, 1996)
(AMNH 5434; = AMNH 5336 of Matthew and Brown, 1923 and Russell, 1970) (adult) skull (1.05 m), lower jaw (1.025 m), cervical vertebrae, dorsal vertebrae, 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), (femur ~1.093 m) (Matthew and Brown 1923)
(AMNH 5458) (8.6 m, 2.5 tons, adult) skull (990 mm), mandible (985 mm), 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, Carr 1999)
(AMNH 5623)
(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 1923)
(CMN 350) femur (930 mm), tibia (842 norm, 882 mm with astr)
(CMN 2193) (adult) surangular (Carr 1999)
(CMN 2250) ilium (Ford and Chure, 2001)
(CMN 2270) (juvenile) maxilla (Carr 1996, 1999)
(CMN 8782) fragmentary skull, partial skeleton
(CMN 11593) proximal tail, pelvis, hindlimbs including femur (940 mm), tibia (900 mm), metatarsus (580 mm)
(CMN 12063) (juvenile) maxilla (Carr 1996, 1999)
(FMNH PR2211) (130 kg; 5 year old juvenile) postcranial skeleton including ribs, gastralia, femur (445 mm), and fibula
(MOR 557) supraoccipital
(ROM 436) (juvenile) partial premaxilla, maxillary fragment (Carr 1996, 1999)
(ROM 672) metacarpal I, metacarpal II, metacarpal III
(ROM 683) premaxillae, incomplete maxillae, nasal fragment, anterior dentary (Carr, 1996)
(ROM 1237) skeleton lacking presacral vertebrae and hindlimb (Ford and Chure, 2001)
(ROM 1246) dentary
(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) (Carr, 1996)
(ROM 1422) (adult) partial skull (premaxilla, incomplete maxillae, incomplete nasals, lacrimal, postorbital, partial squamosal, quadratojugal, quadrates, partial palatine, partial pterygoid), partial surangular, angular (Carr 1996, 1999)
(ROM 3520) frontal
(ROM 4591) (adult) nasals (Carr 1999)
(RTMP 67.9.164) dentary (445 mm)
(RTMP 67.14.3) frontal
(RTMP 68.3.1) pelvis, hindlimbs
(RTMP 73.30.1) (750 kg; 14 year old subadult) proximal tail, pelvis, hindlimb including tibia (805 mm), metatarsus (515 mm) (femur ~804 mm)
(RTMP 80.16.485) (juvenile) frontal
(RTMP 81.39.8) frontal
(RTMP 82.16.181) frontal
(RTMP 82.28.1) dentary
(RTMP 83.36.100) (juvenile) skull (Carr 1999)
(RTMP 83.36.134) dentary
(RTMP 85.11.3) (juvenile) maxilla (Currie, 1990; Carr 1999)
(RTMP 85.62.1)
(RTMP 86.49.29) dentary
(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) almost complete skeleton including skull (670 mm), mandible and furcula (172 mm) (Makovicky and Currie 1998, Carr 1999)
(RTMP 91.36.533) frontal
(RTMP 91.163.1) skull, skeleton
(RTMP 92.36.76) frontal
(RTMP 92.36.82) squamosal
(RTMP 92.36.749) dentary
(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, furcula (225 mm), femur (916 mm) (Makovicky and Currie 1998)
(RTMP 95.5.1) skull, dentary, skeleton
(RTMP 99.33.1) (607 kg; 14 year old subadult) skull, dentary, skeleton including femur (750 mm)
(RTMP 99.55.170) dentary
(RTMP 2000.12.11) skull
(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)
(UA 10) (adult) skull (900 mm), dentary, several presacral vertebrae, ribs, humerus, metatarsus (Carr 1999)
material (Tokaryk, 1988; Tokaryk and Harington, 1992)
Late Campanian, Late Cretaceous
Judith River Group, Montana, US
(AMNH 3963) (21 years old) dentary, premaxillary tooth (Cope, 1876)
(Peebles coll.) incomplete skeleton (Anonymous, 1995)
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- This entry is incomplete.
Tokaryk (1988) and Tokaryk and Harington (1992) reported material from The Dinosaur Park Formation of Saskatchewan.
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.
At the Armour Symposium (2001), Currie reported skin impressions associated with the holotype of Gorgosaurus, which lacked scales. 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 has also seen tyrannosaurid skin impressions at the MOR.
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.
Carr, 1998. Tyrannosaurid (Dinosauria: Theropoda) craniofacial ontogeny: comparative parsimony analysis of ontogenetic characters. JVP 18(3) 31A
Mackovicky, P. J. and Currie, P. J., 1998. "The presence of a furcula in tyrannosaurid theropods, and its phylogenetic and functional implications," Journal of Vertebrate Paleontology 18(1): 143-149 [April 10, 1998].
http://www.cmnh.org/dinoarch/1995Mar/msg00410.html
http://www.cmnh.org/dinoarch/1996Feb/msg00497.html
Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology.19: 497-520.
http://www.cmnh.org/dinoarch/2001May/msg00383.html
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 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 (Holtz, 2001), 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 (Smith, 2005). 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.
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. Contrib. Canad. Pal. III. 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.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. Tanke and Carpenter (eds). Mesozoic Vertebrate Life. Indiana University Press, Bloomington. pp 64-83.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Smith, 2005. Heterodonty in Tyrannosaurus rex: Implications for the taxonomic and systematic utility of theropod dentitions. Journal of Vertebrate Paleontology. 25(4), 865-887.

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 (dentary 905 mm)
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), 3 pedal unguals (109 mm)
Referred- (AMNH 5218 small individual) (~4.1 m) pedal phalanx III-3 (42 mm) (Currie, 2000)
?(AMNH 5218 large individual; may be Daspletosaurus) (~8.7 m) pedal phalanx III-3 (99 mm) (Currie, 2000)
(AMNH 5218; part of bonebed containing AMNH 5218-5235) two dentaries, fourteen vertebrae, two chevrons, scapula, coracoid, pubes, femur, two tibae, partial fibula, astragalus, calcaneum, nine metatarsals, sixteen pedal phalanges, seven pedal unguals (Currie, 2000)
(AMNH 5222) incomplete skull (Carr, 1999)
(AMNH 5224) (several individuals) includes skull, tail and hindlimbs (Ford and Chure, 2001)
(AMNH 5228) (~6.9 m) metatarsal III, metatarsal IV (465 mm) (Currie, 2000)
....(AMNH 5218) pedal phalanx II-2, phalanx IV-3 (Currie, 2000)
(AMNH 5229) (~5.1 m) metatarsal IV (394 mm) (Currie, 2000)
....(AMNH 5218) pedal phalanx III-2, phalanx III-3 (Currie, 2000)
(AMNH 5230; lost) metatarsus (Currie, 2000)
(AMNH 5231) (~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 5218) femur, tibia, phalanx II-1, phalanx II-2, phalanx III-1, phalanx IV-1, phalanx IV-2 (Currie, 2000)
(AMNH 5232) (~8.1 m) two distal tarsals, metatarsal II, metatarsal III, metatarsal IV (521 mm), metatarsal V (Currie, 2000)
....(AMNH 5218) pedal phalanx I-1, phalanx III-1, phalanx IV-1 (Currie, 2000)
....(AMNH 5226) 25 caudal vertebrae (Currie, 2000)
....(AMNH 5227) tibia, fibula, astragalus (Currie, 2000)
(AMNH 5233) (~6.0 m) metatarsal II, phalanx II-1, metatarsal III, phalanx III-1, metatarsal IV(426 mm) (Currie, 2000)
....(AMNH 5218) pedal phalanx I-1, phalanx III-3, phalanx IV-1, phalanx IV-3, phalanx IV-4 (Currie, 2000)
(AMNH 5234) (~6.8 m) astragalus, metatarsal II, metatarsal III, phalanx III-1, metatarsal IV (452 mm), phalanx IV-1 (Currie, 2000)
....(AMNH 5218) humerus, pedal phalanx II-1, phalanx III-2, phalanx III-3, phalanx IV-3 (Currie, 2000)
(AMNH 5235) (~7.3 m) femur, metatarsal II, metatarsal III, metatarsal IV (486 mm), ungual (Currie, 2000)
....(AMNH 5218) humerus, pedal phalanx I-1, phalanx II-1, phalanx III-2 (Currie, 2000)
?(AMNH 5255) hindlimb (Osborn, 1916)
(CMN 2196; see also CMN 2196 under Dinosaur Park Fm. indet.) gastralia, scapulocoracoid (www.paleofile.com)
(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)
(RTMP 81.10.1) (8 m; 1.14 tons; 24 year old adult) partial skull (970 mm) (maxilla, jugal, ectopterygoid, quadrates) surangular, angular, prearticular, partial postcrania missing tail (femur- 895mm, tibia- 970 mm, metatarsus- 610 mm) (Paul, 1988)
(RTMP 85.98.1) (subadult) incomplete skull, partial skeleton (Carr, 1999)
(RTMP 86.64.1) (6.5 m; 760 kg; 15 year old subadult) incomplete skeleton including skull, furcula (192 mm) and femur (~782 mm) (Carr, 1999)
(RTMP 86.205.1) quadrate, basioccipital, braincase elements, cranial fragments, mandibles, ribs, forelimb and hindlimb elements (Currie, 2003)
(RTMP 94.186.1) fragmentary skeleton, skin impressions (Currie, 2003)
(RTMP 97.58.1) skull, skeleton (Currie, 2003)
material (Evans et al., 2003)
Late Cretaceous
Alberta
(RTMP coll.) caudal series, tibiae, fibulae, pes (Rondeau, 1995)
localities not yet entered
(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) (Makovicky and Currie 1998, Carr 1999)
(ROM 12790) skull (Carr 1999)
(RTMP 81.9.1) frontal
?(RTMP 81.31.59) tooth (Erickson, 1995)
(RTMP 82.13.3) postorbital
(RTMP 94.25.6) dentary
(RTMP 95.25.83) maxilla
(RTMP 98.63.87) vomer
(RTMP 98.63.88) maxilla
(RTMP 99.50.140) maxilla
(RTMP 2002.45.46) (50.3 kg; 2 year old juvenile) (femur ~316 mm)
?(RTMP coll.) twenty-five teeth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 2000)
dentary, teeth (Ryan, Bell and Eberth, 1995)
Diagnosis- (after Carr, 2005) interfenestral strut narrow; medial frontal process of nasal elongate; paired medial frontal processes are lanceolate; lateral frontal processes of nasal are short; dorsal surface of lateral frontal process is convex; ventral margin of the joint surface for the quadratojugal on the jugal extends anterodorsally at a steep angle; dorsal ridge of postorbital boss positioned posterior to the boss; dorsotemporal fossa extends on to the lateral surface of the squamosal as a lip-like ridge; anteriorly directed flange present on distal end of the dorsal quadratojugal process; posterior pneumatic foremon of the palatine is large; the joint surface for the fibula is on the ventrolateral surface of the lateral cnemial process of the tibia.
(from Currie et al., 2003) most specimens of have more numerous, deeper pits in the ventral surfaces of the maxillary palatal shelves to accommodate the tip of the dentary teeth; the occipital condyle is oriented more ventrally than in Gorgosaurus, although not to the same degree as in the tyrannosaurines; the braincase box (Bakker et al.1988) is mediolaterally wider than anteroposteriorly long, in contrast with Gorgosaurus where the dimensions are the opposite; the prefrontal seems to have very limited dorsal exposure; the lacrimal did not plug into a socket in the frontal, which is more similar to Tyrannosaurus than Gorgosaurus; an angular suture between the exoccipital and basioccipital in the occipital condyle.
(from Carr, 1996) less maxillary teeth (12 vs. 13-15); antorbital fossa not emarginated dorsally below maxillary fenestra; robust basicranium for size (individual variation?); foramen set within deep sulcus in basioccipital (individual variation?); perforation present at ventral border of pneumatic recess that excavates basioccipital ventrolaterally (postmortem damage?).
Comments- This entry is incomplete.
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.
References- Cope, 1892.
Lambe, 1904.
Osborn, 1916. Skeletal adaptation of Ornitholestes, Struthiomimus, Tyrannosaurus. Bulletin of the American Museum of Natural History. 35, 733-771.
Mackovicky 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: 497-520.
Evans, Lam, Maddin and Conacher, 2003.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
A? sp. indet. (Buckley et al., 2005)
Late Campanian-Early Maastrichtian, Late Cretaceous
Wapiti Formation, British Columbia, Canada
Material- partial tooth
Reference- Buckley, McCrea and Currie, 2005.

Tyrannosaurinae Osborn, 1906 sensu Matthew and Brown, 1922
Definition- (Tyrannosaurus rex <- Gorgosaurus libratus, Albertosaurus sarcophagus) (Sereno, in prep.)
Other definitions- (Tyrannosaurus rex <- Albertosaurus sarcophagus, Daspletosaurus torosus, Gorgosaurus libratus) (modified from Sereno, 1998)
(Tyrannosaurus rex <- Aublysodon mirandus) (modified from Holtz, 2001)
(Tyrannosaurus rex <- Albertosaurus sarcophagus) (Holtz, 2004; modified from Currie et al., 2003)
= Shanshanosaurinae Dong, 1977 sensu Olshevsky, 1995
= Tyrannosaurinae sensu Currie et al., 2003
Definition- (Tyrannosaurus rex <- Albertosaurus sarcophagus) (modified)
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- Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.

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)
Comments- This taxon was originally named and described by Carr in his unpublished thesis (Carr, 2005) before being officially described by Carr et al. (2011). It was first mentioned by Stadtman et al. (1999) as two individuals, but is actually from 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.
References- Stadtman, Chure, Scheetz and Britt, 1999. Fossil vertebrates from the Kaiparowitz Fm. (Late Cretaceous), Grand Staircase-Escalante Monument (GRST), Utah. JVP. 19(3), 77A.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
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.

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- Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. 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- Carr, 2005) to clade with OMNH 10131 (a juvenile specimen described as Aublysodon but now referred to an unnamed albertosaurine- Carr, 2005). 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 tyrannosaurids (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 (Molnar and Carpenter, 1989), 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, New York.
Molnar and Carpenter, 1989. The Jordan theropod (Maastrichtian, Montana, U.S.A.) referred to the genus Aublysodon. Geobios. 22, 445-454.
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, Cambridge. pp 434-455.
Holtz, 2001. The phylogeny and taxonomy of the Tyrannosauridae. Tanke and Carpenter (eds). Mesozoic Vertebrate Life. Indiana University Press, Bloomington. pp 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.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
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 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 (Currie, 2003) and Tyrannosaurus (LACM 28471), while vertical ridges are present in Gorgosaurus and juvenile Tyrannosaurus as well (Carr and Williamson, 2004). 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. Proc Acad. Nat. Sci. 8(2), 72.
Cope, 1866. Discovery of a gigantic dinosaur in the Cretaceous of New Jersey Proc. Acad. Nat. Sci. Philadelphia. 18, 275-279.
Leidy, 1868. Remarks on a jaw fragment of Megalosaurus. Proc. Acad. Nat Sci. 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.
Hay, 1930. Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution of Washington. 390(II), 1-1074.
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, 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.
Currie, 2005. Theropods, including birds. In Dinosaur Provincial Park, a Spectacular Ancient Ecosystem Revealed. Currie and Koppelhus (eds). Indiana University Press, Bloomington, Indiana. 367-397.

Daspletosaurus Russell, 1970
Diagnosis- (after Carr, 2005) lateral surface of maxilla anterior to antorbital fenestra is coarse; anteromedial lacrimal process reaches dorsal margin of antorbital fossa in lateral view; hornlet is present on the lateral surface of the posterior lacrimal process; jugal horn broad in ventral view; in lateral view the anterodorsal squamosal process stops posterior to the level of the anterior margin of the laterotemporal fenestra; orbital margin of postorbital vertical.
References- Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
D. sp. nov. (Varricchio and Currie, 1991)
Mid-Late Campanian, Late Cretaceous
Upper Two Medicine Farmation, Montana, US
Material- (MOR 590) (adult) skull, mandible, hindlimb (Carr, 1999)
?(RMDRC coll.; Pete) (11 m) 70% complete skeleton including ribs (RMDRC online)
(OTM 200) dentary, splenial, several teeth, cervical vertebrae, dorsal vertebrae, ribs, sacral vertebrae, caudal vertebrae, chevrons, partial ilia, pubes, ischium (Varricchio, 2001)
(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)
skull, hindlimb (Varricchio and Currie, 1991)
Diagnosis- (after Carr, 2005) dorsal process of palatine extends vertically.
Comments- Horner et al. (1992) thought this taxon 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. Carr (2005) recovered it as the sister taxon of Daspletosaurus torosus and an undescribed Daspletosaurus species from the Dinosaur Park Formation, where it is placed here.
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.
References- Varricchio and Currie, 1991. New theropod finds from the Two Medicine Formation (Campanian) of Montana. Journal of Vertebrate Paleontology. 12 (suppl. to no. 3): 59A.
Horner and Varricchio, 1992.
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: 497-520.
Varricchio, 1999.
Varricchio, 2001. Gut contents from a Cretaceous tyrannosaurid: Implications for theropod dinosaur digestive tracts. Journal of Paleontology. 75 (2): 401-406.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
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: The Carnivorous Dinosaurs, Edited by Carpenter, K., III. Theropods as living animals, p. 313-324.
Daspletosaurus Russell, 1970 sensu Paul, 1988
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 an undescribed species from the Two Medicine Formation. On this website, Paul's (1988) subgenus Daspletosaurus is used as a label for it.
Diagnosis- (after Carr, 2005) postorbital boss approaches laterotemporal fenestra.
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)
(RTMP 2001.36.1) skull, skeleton (Currie, 2003)
Comments- Although many other specimens are usually referred to this species, Currie (2003) has noted those from the Dinosaur Park Formation belong to a new undescribed species. 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: Fourth Symposium on Mesozoic Terrestrial Ecosystems, short papers, edited by Currie, P. J., and Koster, E. H., p. 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, v. 50, n. 1, p. 93-99.
D. (D.) sp. nov. (Currie et al., 2003)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Material- (AMNH 5438) (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)
(BMNH R4863) premaxilla, maxilla, dentary, hyoid (Russell, 1970)
(CMN 841) (adult) incomplete postorbital (Carr 1996, 1999)
(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)
(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 (Carr, 1999)
(RTMP 92.36.1220) skull, skeleton (Carr, 1999)
(RTMP 94.143.1) (5.8 m; 496 kg; 10 year old subadult) skull (620 mm), 10+ vertebrae, ribs, fractured ilium (femur ~626 mm) (Tanke and Currie, 2000)
(RTMP 94.218.1) (juvenile) skull (Carr, 1999)
(RTMP coll.) skull (Currie and Russell, 2005)
(UA 11) femur (1 m), metatarsal IV(490 mm) (Russell, 1970)
material (Ryan et al., 2001)
Comments- Currie (2003) notes specimens of Daspletosaurus from the Dinosaur Park Formation appear to represent a distinct species from the holotype, citing a paper in preperation by Currie and Bakker.
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.
References- Russell, 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National Museum of Natural Science Publications in Palaeontology. 1: 1–34.
Carr, 1998. Tyrannosaurid (Dinosauria: Theropoda) craniofacial ontogeny: comparative parsimony analysis of ontogenetic characters. JVP 18(3) 31A
Mackovicky 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: 497-520.
Tanke and Currie, 2000. Head-biting behavior in theropod dinosaurs: paleobathological evidence. Gaia 15. 167-184.
Carr and Williamson, 2001. Resolving tyrannosaurid diversity: Skeletal remains referred to Aublysodon belong to Tyrannosaurus rex and Daspletosaurus. JVP 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, v. 16, p. 482-506.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Currie, 2005. Theropods, including birds: In: Dinosaur Provincial Park, a spectacular ecosystem revealed, Part Two, Flora and Fauna from the park, Chapter 19, edited by Currie, P. J., and Koppelhus, E. B., Indiana University Press, p. 367-397.
Currie and Russell, 2005. The geographic and stratigraphic distribution of articulated and associated dinosaur remains. In: Dinosaur Provincial Park, a spectacular ecosystem revealed, Part Three, Interpretations, Chapter 28, edited by Currie, P. J., and Koppelhus, E. B., Indiana University Press, p. 537-569.
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 89.17.53) maxilla
(RTMP 91.36.403) frontal
(RTMP 94.172.115) maxilla
(RTMP 98.48.1) maxilla, nasal
(SDNH 32701) frontal
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).
Reference- Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology.19: 497-520.
D? sp. (Langston, Standhardt and Stevens, 1989)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Comments- These include remains from both the upper and lower sections of the formation.
Reference- Langston, Standhardt and Stevens, 1989. Fossil vertebrate collecting in the Big Bend – history and perspective, p. 11-21. In Busbey, A.B. III, and T. M. Lehman, (eds.). Vertebrate paleontology, biostratigraphy, and depositional environments, Latest Cretaceous and Tertiary, Big Bend area, Texas. Guidebook, 49th annual meeting of the Society of Vertebrate Paleontology, Austin, Texas.
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 a new genus by Carr (2005). The present specimen may belong to this new genus as well.
References- Carr and Williamson, 2000. A review of Trannosauridae (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. Unpublished PhD dissertation. 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: Late Cretaceous Vertebrates from the Western Interior, edited by Lucas, S. G., and Sullivan, R. M., New Mexico Museum of Natural History & Science, Bulletin 35, p. 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 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 an undescribed taxon (Carr, 2005). It's quite possible the present specimens belong to this new genus as well, though they may be too fragmentary to assign to any genus.
References- Carr and Williamson, 2000. A review of Trannosauridae (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. Unpublished PhD dissertation. 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.

Tyrannosaurinae sensu Sereno, 1998
Definition- (Tyrannosaurus rex <- Albertosaurus sarcophagus, Daspletosaurus torosus, Gorgosaurus libratus) (modified)
Diagnosis- (after Carr, 2005) maxillary fenestra extends anteromedial to the anterior margin of the antorbital fossa; antorbital fossa reaches maxillonasal suture with elongate contact; accessory pneumatic foramen in anterior lacrimal process is distal in position; joint surface for the quadratojugal on the jugal extends anteriorly from the ventral jugal margin; posterodorsal jugal process extends posterodorsally; lingual bar of the dentary flanks anterior two alveoli; oval scar of the femur is on the posteromedial edge of the bone; the indentation of the lateral cnemial process of the tibia is anterior to the midlength of the process; the posteroventral heel of the calcaneum is short or absent.
Comments- This clade is called Tyrannosaurus by several authors (Carpenter, 1992; Holtz, 1994, 1995, 2001; Carr, 1999, 2005; Carr et al., 2005), while others (Currie, 2003; Hurum and Sabath, 2003; Holtz, 2004) retain bataar and rex in separate genera.
References- Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.

Alioramini Olshevsky, 1995
Alioramus Kurzanov, 1976
A. remotus Kurzanov, 1976
= Alioramus altai Brusatte, Carr, Erickson, Bever and Norell, 2009
Maastrichtian, Late Cretaceous
Nogonn Tsav Beds, Mongolia
Holotype- (GI 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
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)
Diagnosis- (after Kurzanov, 1976) elongate skull (lengfth/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- 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 holotype. 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). 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; postorbital lacks an orbital process; more dentary teeth; muscular fossa above surangular foramen faces mostly dorsally; laterally projecting jugal horn; 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. Whle 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). The fact most differences could be explained by ontogeny, coupled with the unique similarities present in Alioramus and the contemporaneous Tarbosaurus strongly suggest the former is a juvenile of the latter. The alternative presented by Brusatte et al., where a distinct genus known only from juvenile specimens and based on characters largely found in juvenile tyrannosaurs is contemporaneous with a taxon it is not the sister group of yet shares autapomorphies with, is considered unlikely.
Brusatte et al. (2009) erected a new species Alioramus altai based on a partial skeleton discovered in 2001 from the contemporaneous Nemegt Formation. 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.
References- Kurzanov, 1976. A new Late Cretaceous carnosaur from Nogon-Tsav Mongolia. Sovmestnaa Sovetsko-Mongolskaa Paleontologiceskaa Ekspeditcia, Trudy. 3, 93-104.
Currie, 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologica Polonica. 48(2), 191-226.
Currie, Hurum and Sabath, 2003. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontologica Polonica. 48(2), 227-234.
Hurum and Sabath, 2003. Giant theropod dinosaurs from Asia and North America: Skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologica Polonica. 48(2), 161-190.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds). The Dinosauria Second Edition. University of California Press. 861 pp.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
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.
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.

Tarbosaurini Olshevsky, 1995
Tarbosaurus Maleev, 1955
= Shanshanosaurus Dong, 1977
= Maleevosaurus Carpenter, 1992
= Jenghizkhan Olshevsky, 1995
T. bataar (Maleev, 1955) Rozhdestvensky, 1965
= Tyrannosaurus bataar Maleev, 1955
= Gorgosaurus novojilovi Maleev, 1955
= Tarbosaurus efremovi Maleev, 1955
= Gorgosaurus lancinator Maleev, 1955
= Deinodon novojilovi (Maleev, 1955) Maleev, 1964
= Deinodon lancinator (Maleev, 1955) Kuhn, 1965
= Aublysodon lancinator (Maleev, 1955) Charig, 1967
= Aublysodon novojilovi (Maleev, 1955) Charig, 1967
= Shanshanosaurus huoyanshanensis Dong, 1977
= Tyrannosaurus efremovi (Maleev, 1955) Rozhdestvensky, 1977
pr= Tyrannosaurus "turpanensis" Zhai, Zhang and Tong, 1978
= Tarbosaurus novojilovi (Maleev, 1955) Olshevsky, 1978
pr= Tyrannosaurus luanchuanensis Dong, 1979
= Aublysodon huoyanshanensis (Dong, 1977) Paul, 1988
= Albertosaurus novojilovi (Maleev, 1955) Mader and Bradley, 1989
pr= Tarbosaurus "turpanensis" (Zhai, Zhang and Tong, 1978) Olshevsky, 1991
pr= Tarbosaurus luanchuanensis (Dong, 1979) Olshevsky, 1991
= Maleevosaurus novojilovi (Maleev, 1955) Carpenter, 1992
= Jenghizkhan bataar (Maleev, 1955) Olshevsky, 1995
= Jenghizkhan luanchuanensis (Dong, 1979) Olshevsky, 1995
= Tyrannosaurus novojilovi (Maleev, 1955) Glut, 1997
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (PIN 551-1) (~12.4 m, ~5 tons) partial skull (~1.35 m), dentary, posterior cervical vertebrae, first four dorsal vertebrae (femur ~1.2 m)
Referred- (930928 NG WTB) cranial fragments (Watabe and Suzuki, 2000)
(940824 BgT TSGT) complete postcranial skeleton (Watabe and Suzuki, 2000)
(940826 BgT OTGN) mandible, postcrania (Watabe and Suzuki, 2000)
(950622 BgT Tarbo. A) caudal vertebrae (Suzuki and Watabe, 2000)
(950622 BgT Tarbo. PJ1-9) elements (Suzuki and Watabe, 2000)
(950626 BgT Tarbo. C) pelvis (Suzuki and Watabe, 2000)
(950817 BgT Tarbo. D) ribs (Suzuki and Watabe, 2000)
(950817 BgT Tarbo. E) gastralia (Suzuki and Watabe, 2000)
(950817 BgT Tarbo. F) gastralia (Suzuki and Watabe, 2000)
(970716-18 KmT) skull, humerus, pelvis, femur, other elements (Watabe and Suzuki, 2000)
(980803 BgT NAR) fragmentary elements (Suzuki and Watabe, 2000)
(IGM 100/59) skull (1.07 m), incomplete 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 100/65) partial skull, surangular (Hurum and Sabath, 2003)
(IGM 100/67) fragmentary skull, braincase (Hurum and Sabath, 2003)
(IGM 100/69) occiput (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) (Currie, 2003; Hurum and Sabath, 2003)
(IGM 107/3) skull (Maleev, 1974; Hurum and Sabath, 2003)
....(PIN 552-1) partial skeleton including femur (970 mm), tibia (870 mm), metatarsal II (455 mm), metatarsal III (540 mm), metatarsal IV (510 mm) (Maleev, 1974)
(IGM 107/7; ?=MPD 107/6A) (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, skin impressions (Tsuihiji et al., 2011)
(IGM 107/14) (three individuals; subadults) cranial elements including nasals (293 mm), lacrimal (172 mm), postorbital, squamosal, braincase, postcranial elements (Tsuihiji et al., 2011)
(No. 1) fragmentary skeleton (Gradzinsky, 1970)
(No. 2) incomplete skeleton (Gradzinsky, 1970)
(No. 4) fragmentary skeleton (Gradzinsky, 1970)
(PIN 551-2; holotype of Tarbosaurus efremovi) (adult) skull, mandibles, incomplete skeleton including femur (970 mm), tibia (880 mm), metatarsus (540 mm) (Maleev, 1955)
(PIN 551-3) (7.7 m, 2.1 tons) skull (1.135 m), dentary, femur (970 mm), metatarsus (546 mm) (Maleev, 1974)
(PIN 551-4) partial skeleton (Maleev, 1974)
(PIN 551-6) incomplete scapulocoracoid (Maleev, 1974)
(PIN 551-91) maxillary fragment (Hurum and Sabath, 2003)
(PIN 552-2; holotype of Gorgosaurus novojilovi) (6.18 m, juvenile) skull lacking quadrate, quadratojugal, parietal and braincase (713 mm), anterior dentary, cervical vertebrae 3-10, cervical ribs, dorsal vertebrae 1-13, dorsal ribs, gastralia, sacrum, caudal vertebrae 1-45, chevrons, scapula, coracoid, humerus (143 mm), radius (108 mm), ulna, manus, ilium (682 mm), pubis (507 mm), ischium (390 mm), femur (650 mm), tibia (781 mm), fibula, astragalus, metatarsal II (365 mm), metatarsal III (420 mm), metatarsal IV (395 mm), pes (Maleev, 1955)
(PIN 552-3) incomplete skull, partial skeleton (www.paleofile.com; typo for 552-2?)
(PIN 552-4) partial skeleton (www.paleofile.com)
(PIN 553-1; holotype of Gorgosaurus lancinator) skull, mandibles, vertebrae, metacarpals, metatarsals (Maleev, 1955)
(PIN 553-2) distal caudal vertebrae (Maleev, 1974)
(PIN 553-3) braincase (Saveliev and Alifanov, 2007)
(PIN 555-5) partial skeleton (www.paleofile.com)
(PJC.2000.9) (juvenile) incomplete skeleton lacking caudal vertebrae and most of pelvis (Currie, 2002)
?(PJC.2001.14) proximal scapula (Currie, 2002)
(Tokyo Natural Science Museum coll.) skull, incomplete skeleton (www.paleofile,com)
(ZPAL MgD-I/3) (5.8 m, 760 kg) skull (745 mm), cervical vertebrae, dorsal vertebrae, ribs, gastralia, ten proximal caudal vertebrae, scapulae, coracoids, forelimbs, ilium, pubis, ischium, femora (700 mm), tibiae (700 mm), fibulae, metatarsi (445 mm), pes (Hurum and Sabath, 2003)
(ZPAL MgD-I/4) skull (1.11 m), mandible (dentary 480 mm), 13 sacral and proximal caudal vertebrae, ilium, femur (970 mm), tibia, fibula, metatarsus (555 mm), pes (Hurum and Sabath, 2003)
(ZPAL MgD-I/5) (large) maxilla, quadrate, mandibles, fragments of eleven ribs, fragmentary ilia, fragmentary pubis, ischia, hindlimb, metatarsal (Hurum and Sabath, 2003)
(ZPAL MgD-I/26) fragmentary maxilla, teeth (Hurum and Sabath, 2003)
(ZPAL MgD-I/29) partial skull, mandible, six cervical vertebrae, eleven ribs, sacrum, 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/31) posterior mandible (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 ribs (Hurum and Sabath, 2003)
(ZPAL MgD-I/52) dentary tooth (Hurum and Sabath, 2003)
(ZPAL MgD-I/67) jugal (Hurum and Sabath, 2003)
(ZPAL MgD-I/93) endocranial cast (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)
?(ZPAL MgD-I coll.) astragalus (105 mm) (Osmolska and Roniewicz, 1970)
(very large) two skeletons (Kielan-Jaworwska and Barsbold, 1972)
(small) two incomplete skeletons (Kielan-Jaworwska and Barsbold, 1972)
three incomplete skeletons (Kielan-Jaworwska and Barsbold, 1972)
material (Perle et al., 1994)
premaxillary tooth (Currie, 2001)
frontal (Currie, 2001)
frontals, skin impressions (Currie, 2001)
tooth (Currie, 2001)
metatarsal IV (Currie, 2001)
frontals, basioccipital (Currie, 2003)
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)
Early Maastrichtian, Late Cretaceous
White Beds of Khermeen Tsav, Mongolia
?material (Gradzinski, Kilean-Jaworowska and Maryanska, 1977; Maryanska, 1977)
Early Maastrichtian, Late Cretaceous
Nemegt Svita (=Beds of Bugeen Tsav), Mongolia
?material (Gradzinski, Kilean-Jaworowska and Maryanska, 1977)
Maastrichtian, Late Cretaceous
Subashi Formation, Xinjiang, China
(IVPP V4878; holotype of Shanshanosaurus huoyanshanensis) (2.3 m, 27 kg, juvenile) (skull ~288 mm) premaxilla (lost), maxilla (180 mm), mandible, tooth (14.4 mm), atlantal centrum, axis (22.3 mm), nine incomplete cervical vertebrae (anterior cervical 21.2 mm), cervical postzygapophysis, anterior cervical rib, thirteen incomplete dorsal vertebrae (posterior dorsal 38.7 mm), several dorsal ribs, scapula (138 mm), coracoid, humerus (88.8 mm), distal pubes, femur (279 mm), proximal tibiae (Dong, 1977)
?(IVPP coll.; material of Tyrannosaurus "turpanensis") five teeth, three posterior sacral vertebrae, ilium (Dong, 1977)
Campanian-Early Maastrichtian, Late Cretaceous
Quiba Formation, Henan, China
?(IVPP V4733; holotype of Tyrannosaurus luanchuanensis) five teeth (35.1-36.9 mm), partial vertebra
Campanian-Maastrichtian, Late Cretaceous
Tsagaan Svita, Russia
?material (Bolotsky and Moisyeyenko, 1988; Nessov, 1995)
? (likely Nemegt Formation, Mongolia)
(GI 100/66) (juvenile) nasals, lacrimal (Currie, 2003)
(GI 100/777; 100/177 in Currie and Dong) (juvenile) premaxilla, two maxillae, vomer (Currie and Dong, 2001; Currie, 2003)
(GI 107/1) skull (991 mm), dentary, coronoid, splenial (Hurum and Sabath, 2003)
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.
Comments- A furcula is known (Sabath pers. comm. to Carpenter and Smith, 2001).
One or more species?- The holotype specimen was first named Tyrannosaurus bataar by Maleev (1955), with smaller spcimens subsequently named Tarbosaurus efremovi, Gorgosaurus lancinator and Gorgosaurus novojilovi (Maleev, 1955). Maleev (1964) later transferred the latter two species to Deinodon. Rozhdestvensky (1965) synonymized all four species into Tarbosaurus bataar, while Maleev (1974; edited and published by Rozhdestvensky and Kurzanov) 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 (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 Carpenter [1990] did show Tyrannosaurus varied 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 properties. 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 (1977) was a proximal rib, while the cervical vertebrae are amphicoelous, not procoelous (contra Molnar, 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 (which may be another juvenile Tarbosaurus any way). 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 to Tarbosaurus sp.. Zhai et al. (1978) later listed the nomen nudum Tyrannosaurus "turpanensis", which based on 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 (I'm assuming the faunas are similar).
Fragmentary remains from the Quiba Formation of China were named Tyrannosaurus luanchuanensis (Dong, 1979), later referred to Tarbosaurus (Olshevsky, 1991) and Jenghizkhan (Olshevsky, 1995). Carr and Williamson (2000) noted its teeth have a denticle density like that of Tyrannosaurus, different from Daspletosaurus and albertosaurines. Since Tarbosaurus has the same density as Tyrannosaurus (Hurum and Sabath, 2003), and the Quiba Formation may be contemporaneous with the Nemegt Formation, it is provisionally considered a junior synonym of T. bataar.
Gradzinski et al. (1977) and Maryanska (1977) have referred material from the White Beds of Khermeen Tsav and the Nemegt Svita to Tarbosaurus, but these remains have not been published. The age of the deposits does make the presence of Tarbosaurus bataar plausible.
Jerzykiewicz, Currie, Eberth, Johnston, Koster and Zheng (1993) referred premaxillary and maxillary teeth from the Late Campanian Djadochta Formation of Mongolia to Tarbosaurus sp.. This is slightly earlier than the Nemegt and Subashi Formations, suggesting they are not from T. bataar at least.
Dong (1979) briefly described some unassociated elements from the Yuanpu (=Nanxiong) Formation of China as Tarbosaurus sp.. As with the Djadochta material, they are from Campanian deposits, suggesting they are not T. bataar even if they are Tarbosaurus.
Nessov (1995) referred a femur (N 601/12457) from the Bostobe Formation of Kazakhstan to Tarbosaurus sp. (incorrectly translated by Olshevsky, DML 1996 as an ilium), but Carr (2005) determined it lacks the synapomorphies of Tarbosaurus + Tyrannosaurus and of Alectrosaurus. It seems to be a Beipiaosaurus-grade therizinosaur. Nessov also referred a mandible from Karachek in Kazakhstan to Tarbosaurus aff. bataar. This has not been rigorously evaluated however, and is probably wrong considering the older age of the deposits. Finally, Nessov noted tyrannosaurid remains referred to by Bolotsky and Moisyeyenko (1988) from the Tsagaan Svita of Russia, which he stated were probably Tarbosaurus sp.. This is possible though unproven.
Another specimen often referred to T. bataar (e.g. Molnar et al., 1990) is T? periculosus from the Tsagaan Svita of China (originally Albertosaurus periculosus). Based only on a tooth, generic assignment to any genus is not yet established, but synonymy with T. bataar is possible.
Chingkankousaurus fragilis is sometimes listed as a junior synonym of Tarbosaurus bataar, but is based on an indeterminate anterior dorsal rib fragment (originally misidentified as a scapula) that may indeed be Tarbosaurus, based on its size, but could also be non-theropod (see entry). 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).
References- Maleev, 1955. Gigantic carnivorous dinosaurs from Mongolia [in Russian]. Doklady AN SSSR. 104 (4): 634–637.
Maleev, 1955. New carnivorous dinosaurs from the Upper Cretaceous of Mongolia [in Russian]. Doklady AN SSSR. 104 (5): 779–782.
Maleev, E.A. 1964. Suborder Theropoda. Carnivorous dinosaurs [in Russian]. In Rozhdestvinsky and Tatarinov (eds.). Osnovy paleontologii. Spravocnik dla paleontologov i geologov SSSR. Zemnovodnye, presmykausiesia, pticy, 529-540. Nauka, Moskva.
Kuhn, 1965. Saurischia: Fossilium Catalogus, I: Animalla, Pars 109, p. 1-94.
Maleev, 1965. On the brain of predatory dinosaurs. Paleontol. Zh.. 2, 141-143.
Rozhdestvenskiy, 1965. Growth changes and some problems of systematics of Asian dinosaurs [in Russian]. Paleontologiceskij zurnal. 3: 95–109.
Charig, 1967.
Gradzinski, 1970. Sedimentation of dinosaur-bearing Upper Cretaceous deposits of the Nemegt Basin, Gobi Desert. Palaeontologia Polonica. 21: 147–229.
Osmolska and Roniewicz, 1970. Deinocheiridae, a new family of theropod dinosaurs. Palaeontol. Polonica 21: 5-19.
Kielan-Jaworowska and Barsbold, 1972. Narrative of the Polish-Mongolian Paleontological Expedition 1967–1971. Palaeontologia Polonica. 27: 5–16.
Maleev, 1974. Gigantic carnosaurs of the family Tyrannosauridae. (In Russian]. In: N.N. Kramarenko (ed.), Fauna i biostratigrafia mezozoa i kainozoa Mongolii. Sovmestnaa Sovetsko-Mongolskaa Paleontologiceskaa Ekspedicia, Trudy 1: 132–191.
Dong, 1977. On the dinosaurian remains from Turpan, Xinjiang. Vertebrata PalAsiatica 15 (1): 59–66.
Gradzinski, Kielan-Jaworowska and Maryanska, 1977. Upper Cretaceous Djadokhta, Barun Goyot and Nemegt formations of Mongolia, including remarks on previous subdivisions. Acta Geologica Polonica. 27: 281–318.
Maryanska, 1977. Ankylosauridae (Dinosauria) from Mongolia. Paleontol. Polonica 37, 85-151.
Rozhdestvensky, 1977. The study of dinosaurs in Asia. J. Palaeontol. Soc. India 20:102-119.
Olshevsky, 1978. The Archosaurian Taxa (excluding the Crocodylia). Mesozoic Meanderings 1: 1-50.
Zhai, Zheng and Tong, 1978. Stratigraphy of the mammal-bearing Tertiary of the Turfan Basin, Sinkiang. Memoirs of the Institute of Vertebrate Paleontology and Paleoantropology 13: 68–81.
Dong, 1979. [The Cretaceous dinosaur fossils in southern China]. In: Mesozoic and Cenozoic Red Beds in Southern China. Inst. Vert. Paleontol. Paleoanthropol. Nanjing Geol. Paleontol. Inst. Sci.. Press, Beijing. Pp.342-350.
Dong, 1979. Cretaceous Dinosaurs of Hunan, in Mesozoic-Cenozoic Redbeds of Hunan. Palaeontologica Sinica, Pp.346-347. (in Chinese)
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Joint Sovjet-Mongolian Paleontol. Expedition Trans. 19, 5-120 [In Russian].
Bolotsky and Moiseenko, 1988. On Pre-Amurian Dinosaurs. Akad. Nauk. SSSR, Dal'nevostkhnoe Otdelenie Amursky Kompleksny Naukno-Issledovatelsky Institut, Blagoveshensk, Pp. 1-37.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Mader and Bradley, 1989. A redescription and revised diagnosis of the syntypes of the Mongolian tyrannosaur Alectrosaurus olseni. Journal of Vertebrate Paleontology 9 (1): 41–55.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In: Dinosauria, edited by Weishampel, D. B., Dodson, P., and Osmolska, H., California University Press, p. 169-209.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings #2 (1st printing): iv + 196 pp.
Carpenter, 1992. Tyrannosaurids (Dinosauria) of Asia and North America. In: Mateer N, Chen PJ, eds. Aspects of nonmarine Cretaceous geology. Beijing, China: Ocean Press, 250–268.
Jerzykiewicz, Currie, Eberth, Johnston, Koster and Zheng, 1993. Djadoktha Formation correlative strata in Chinese Inner Mongolia: an overview of the stratigraphy sedimentary geology, and paleontology and comparisons with the type locality in the pre-Altai Gobi. Canadian Journal of Earth Sciences 30: 2180-2195.
Perle, Chiappe, Rinchen, Clark and Norell, 1994. Skeletal Morphology of Mononykus olecranus (Theropoda: Avialae) from the Late Cretaceous of Mongolia. American Museum Novitates 3105:1-29.
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.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids [in Japanese]. Kyoryugaku Saizensen (Dino Frontline) 9: 92–119; 10: 75–99.
Glut, 1997. Dinosaurs, the Encyclopedia: Mcfarland & Company, Inc., Publishers, 1076 pp.
Carr, 1998. Tyrannosaurid (Dinosauria: Theropoda) craniofacial ontogeny: comparative parsimony analysis of ontogenetic characters. JVP 18(3) 31A.
Carr and Williamson, 2000. A review of Trannosauridae (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.
Hurum and Currie, 2000. The crushing bite in tyrannosaurids. Journal of Vertebrate Paleontology. 20: 619–621.
Suzuki and Watabe, 2000.
Watabe and Suzuki, 2000. Cretaceous fossil localities and a list of fossils collected by the Hayashibara Museum of Natural Sciences and Mongolian Paleontological Center Joint Paleontological Expedition (JMJPE) from 1993 through 1998. Hayashibara Museum of Natural Sciences Research Bulletin 1: 99–108.
Currie, 2001. Nomadic Expediations, Inc., report of fieldwork in Mongolia, September 2000. Alberta Paleontological Society, Fifth Annual Symposium, Abstract Volume. 8-12.
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: 1729–1737.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. Alberta Paleontological Society, Sixth Annual Symposium, "Fossils 2002", Abstract Volume. 8-12.
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.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
Saveliev and Alifanov, 2007. A new study of the brain of the predatory dinosaur Tarbosaurus bataar (Theropoda, Tyrannosauridae). Paleontological Journal. 41(3), 281-289.
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.
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.
T? sp. indet. (Gilmore, 1933)
Late Campanian, Late Cretaceous
Djadochta Formation, Inner Mongolia, China
Material- ?(AMNH 6522) (~8 m) partial ilium (Gilmore 1933)
premaxillary teeth, maxillary teeth (Jerzykiewicz et al., 1993)
Comments- The teeth were referred to Tarbosaurus sp.. by Jerzykiewicz, which is possible though they are too early to be from T. bataar. Gilmore (1933) described the ilium as a large theropod, perhaps a 'deinodontid'. It is identified as a tyrannosaurid on the AMNH website. They are more likely to be the contemporaneous Zhuchengotyrannus.
References- Gilmore, 1933. Two new dinosaurian reptiles from Mongolia with notes on some fragmentary specimens. American Museum Novitates. 679, 1-20.
http://paleo.amnh.org/fossil/show.html?cat_num=FR%206522
T? sp. indet. (Dong, 1979)
Campanian, Late Cretaceous
Yuanpu (=Nanxiong) Formation, Guandong, China
Material- (IVPP coll.; unassociated) third premaxillary tooth, lateral tooth (72 mm), dorsal vertebra, fragmentary pedal elements
Comments- Dong referred these remains to Tarbosaurus sp.. As these are from earlier deposits than the Maastrichtian Nemegt and Subashi Formations, they are probably not from Tarbosaurus bataar, and may not even belong to this genus. Their serration density is similar to Tarbosaurus and Tyrannosaurus, but not unequivocally different from large Campanian North American teeth. They are more likely to be the contemporaneous Zhuchengotyrannus.
Reference- Dong, 1979. Cretaceous dinosaurs of Hunan, China. Mesozoic and Cenozoic Red Beds of South China: Selected Papers from the "Cretaceous-Tertiary Workshop". Institute of Vertebrate Paleontology and Paleoanthropology & Nanjing Institute of Paleontology (eds.). Science Press, Nanxiong, China. 342-350.

unnamed tyrannosaurine (Khozatsky, 1957)
Santonian-Early Campanian, Late Cretaceous
Kara-Cheku, Almaty, Kazakhstan
Material- (IZK 33/MP-61) incomplete dentary
Comments- This specimen was discovered in 1950 and originally referred to Tyrannosaurus aff. bataar by Khozatsky (1957) and Bazhanov and Kostenko (1958), and later to Tarbosaurus aff. bataar by Nessov (1995). Averianov et al. (2012) redescribed it, finding the reduced first dentary alveolus to place it in the clade of derived tyrannosaurines including Tarbosaurus, Zhuchengtyrannus and Tyrannosaurus. It differs from these taxa in lacking a rugose symphysis.
References- Khozatsky, 1957. [To the history of trionychid turtles in Kazakhstan]. Izvestiya Akademii Nauk Kazakhskoi SSR, Seriya Biologicheskaya. 2, 15-30.
Bazhanov and Kostenko, 1958. [Scheme of stratigraphy of Tertiary deposits of South-Eastern Kazakhstan and Northern Kirghizia in light of paleontological data]. Materialy po Istorii Fauny i Flory Kazakhstana. 2, 5-16.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov, ekologii i paleobiogeografii [Dinosaurs of northern Eurasia: new data about assemblages, ecology, and paleobiogeography]. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Averianov, Sues and Tleuberdina, 2012. The forgotten dinosaurs of Zhetysu (Eastern Kazakhstan; Late Cretaceous). Proceedings of the Zoological Institute RAS. 316(2), 139-147.

Tyrannosaurus? zhuchengensis Hu, Cheng, Pang and Fang, 2001
Campanian, Late Cretaceous
Upper Xingezhuang Formation, Wangshi Series, Shandong, China
Syntypes- (NGMC V1777) metatarsal II (531 mm)
?(NGMC V286) tooth
?(NGMC V288) (juvenile) tooth
?(NGMC V1174) (juvenile) tooth
?(NGMC V1773) tooth
Comments- Originally referred to cf. Tyrannosaurus rex by Hu (1973) and Dong (1979), this material was named Tyrannosaurus zhuchengensis by Hu et al. (2001) in their Shantungosaurus monograph. Whether a holotype was specified is uncertain, as the description has not been translated from Chinese. There is no evidence the material belongs to one individual or one taxon, and indeed two teeth are from juveniles unlike the other two and the metatarsal. Hone et al. (2011) reidentified it as a metatarsal II instead of metatarsal IV. Hone et al. also state the material is indeterminate, so it cannot be referred to the sympatric Zhuchengtyrannus and undescribed tyrannosaurid (ZCDM V0030 and V0032), though it may belong to either. Though Olshevsky (DML, 2002) called it Tarbosaurus zhuchengensis based on geography, this combination has yet to be published.
References- Hu, 1973. A new hadrosaur from the Cretaceous of Zhucheng, Shantung. Acta Geologica Sinica. 2, 179-202.
Dong, 1979. Cretaceous dinosaurs of Hunan, China. Mesozoic and Cenozoic Red Beds of South China: Selected Papers from the "Cretaceous-Tertiary Workshop". Institute of Vertebrate Paleontology and Paleoanthropology & Nanjing Institute of Paleontology (eds.). Science Press, Nanxiong, China. 342-350.
Hu, Cheng, Pang and Fang, 2001. Shantungosaurus giganteus. ISBN 7-116-03472-2. 139 pp.
Olshevsky, DML 2002. 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.

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.
Reference- Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji and Xu, 2011. A new, large tyrannosaurine theropod from the Upper Cretaceous of China. Cretaceous Research. 32(4), 495-503.

undescribed tyrannosaurine (Stein and Triebold, 2005)
Late Campanian, Late Cretaceous
Upper Judith River Formation, Montana, US
Material- (AMNH 30564) gastralium
....(RMDRC 02-001) (Sir William) (~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 2002, 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 as of 2004 at least.
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.
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. In "The origin, systematics, and paleobiology of Tyrannosauridae”, a symposium hosted jointly by Burpee Museum of Natural History and Northern Illinois University, p. 27-28.

Tyrannosaurini Olshevsky, 1995
Tyrannosaurinae sensu Holtz, 2001
Definition- (Tyrannosaurus rex <- Aublysodon mirandus) (modified)
Tyrannosaurus Osborn, 1905
= Manospondylus Cope, 1892 (nomen oblitum)
= Dynamosaurus Osborn, 1905
?= “Clevelanotyrannus” Bakker, Williams and Currie vide Currie, 1987
?= “Nanotyrannes” Anonymous, 1988
?= Nanotyrannus Bakker, Williams and Currie, 1988
= Stygivenator Olshevsky, 1995
= Dinotyrannus Olshevsky, 1995
T. rex Osborn, 1905
= Manospondylus gigas Cope, 1892 (nomen oblitum)
?= Aublysodon amplus Marsh, 1892
?=Aublysodon cristatus Marsh, 1892
?= Deinodon amplus (Marsh, 1892) Hay, 1902
?= Deinodon cristatus (Marsh, 1892) Hay, 1902
= Dynamosaurus imperiosus Osborn, 1905
?= Tyrannosaurus amplus (Marsh, 1892) Hay, 1930
?= Gorgosaurus lancensis Gilmore, 1946
?= Deinodon lancensis (Gilmore, 1946) Kuhn, 1965
?= Aublysodon lancensis (Gilmore, 1946) Charig, 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)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, North Dakota, South Dakota, Wyoming, US
Holotype- (CMN 9380; =AMNH 973) (12.4 m, 4.7 tons; adult) maxilla (695 mm), lacrimals, squamosal, ectopterygoid, dentaries (860 mm), surangular (610 mm), teeth, ninth cervical vertebra, second dorsal vertebra, eighth dorsal vertebra (130 mm), ninth dorsal vertebra (145 mm), tenth dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal vertebra, thirteenth dorsal vertebra (170 mm), dorsal ribs, three gastralia, sacrum (940 mm), scapula (950 mm), humerus (360 mm), ilia (1.515 m), pubes (1.25 m), ischia (1.11 m), femur (1.28 m), tibia (1.14 m), metatarsal I, metatarsal II (615 mm), distal metatarsal III (~684 mm), metatarsal IV (600 mm), phalanx IV-1
Referred- (AMNH 1011) incomplete tooth (Molnar, 1991)
(AMNH 5005) (juvenile) cranial fragments, femur (not collected), fibula (Molnar, 1991)
(AMNH 5020) metatarsal IV (Molnar, 1991)
(AMNH 5021) pedal phalanx (Molnar, 1991)
(AMNH 5027) (12.4 m, 5.7 tons, adult) skull (1.355 m; maxillae 710 mm), mandibles (1.205 m; dentary 850 mm), cervical vertebrae 1-10 (960 mm total), nine cervical ribs, (dorsal series 2.184 m) dorsal vertebrae (~160 mm), twelfth dorsal vertebra (161 mm), thirteenth dorsal vertebra, twenty dorsal ribs, sacrum, first caudal vertebra, caudal vertebrae 1-15, 17, 21, 22, seven chevrons, ilia (1.515 m), pubes (~1.2 m), ischia (1.236 m) (Osborn, 1912)
(AMNH 5044) caudal vertebrae (Molnar, 1991)
(AMNH 5050) partial dentary (Osborn, 1916)
(BHI 3033; Stan) (12.3 m; 3.7 tons; adult) skull (~1.4 m; maxilla 775 mm), mandibles (1.34 m; dentary 915 mm), thirty-five teeth, ten cervical vertebrae, fourteen cervical ribs, thirteen dorsal vertebrae, twelve dorsal ribs, sacrum (1.06 m), thirty-one caudal vertebrae, twenty-four chevrons, ilia (1.55 m), proximal pubes, proximal ischia, femora (1.31 m), tibiae, fibula, astragali, calcanea, metatarsal II (595 mm), metatarsal III, metatarsal IV (600 mm), eleven pedal phalanges (Larson and Frey, 1992; Larson, 2008)
(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 (Horner, 1994; Larson, pers. comm.)
(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, 2008)
(BHI 6219; 007) premaxillae, maxillae, partial dentary, vertebra, dorsal rib, distal humerus, partial tibia, partial fibula, metatarsal, pedal phalanx (Larson, 2008)
(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, 2008)
(BHI 6249; Steven) two incomplete cranial elements, six incomplete dorsal vertebrae, five dorsal ribs, incomplete femur, phalanx, eggshells (Janke, 1996; Larson, 2008)
?(BHI 6235; referred to lancensis) (juvenile) lacrimal?, jugal, frontal, three teeth (Larson, 1995)
(BHI coll.) (subadult) proximal tibia, fibula (Larson, 1995)
(BHI coll.; Rex B; Triceratops Alley rex) premaxilla, maxilla, nasals, lacrimals, frontals, quadratojugal, quadrate, braincase, ectopterygoid, rib, scapula, coracoid (Larson, 2008)
(BMNH R7995; = AMNH 5881) gastralia, femur, tibiae, fibula?, metatarsal I, metatarsal II, metatarsal IV, pedal phalanges? (Osborn, 1906)
?(BMRP 2002.4.1; Jane; referred to lancensis) (~6.4 m; ~680 kg; 11 year old juvenile) incomplete skull (724 mm; maxilla 470 mm), mandible (dentary 505 mm), teeth (~100 mm), seven cervical vertebrae, cervical ribs, four posterior dorsal vertebrae, dorsal ribs, gastralia, sacrum (500 mm), twenty proximal caudal vertebrae (~130 mm), seventeen chevrons, scapulocoracoid, humerus (280 mm), radius, ulna, partial manus, ilia (720 mm), pubes, ischia, femora (720 mm), tibiae (840 mm), metatarsal II (510 mm), metatarsal IV (513 mm), phalanx III-1 (135 mm), phalanx III-2 (103 mm), pedal phalanges (Henderson, 2005)
?(BMRP coll.; Petey) (~7-7.4 m; 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 (Williams, DML 2008)
(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)
(FMNH PR2081; =BHI 2033; Sue; holotype of Tyrannosaurus "stanwinstonorum") (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)
(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, 2008)
(Great Plains Paleontology coll.; Otto) cervical ribs, dorsal ribs, caudal vertebrae, femora, tibiae, fibula, two metatarsals (Larson, 2008)
(KU coll.) (Larson, 1997 pers. comm. to Ford; www.paleofile.com)
(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 molnaris) (~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, 2008)
(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, 2008)
(MOR 555; Wankel rex) (12.4 m; 4.0 tons; adult) incomplete skull (maxilla 798 mm), dentary (990 mm), cervical vertebrae 2-10, dorsal vertebrae 1-13, sacrum (1.01 m), caudal vertebrae 1-18, scapulae (980 mm), coracoids, humerus (377 mm), radius, ulna (198 mm), radiale, ulnare, 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 III, metatarsal IV (605 mm), pedal phalanges (Horner and Lessem, 1993)
(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)
(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, 1982)
(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 (Jacobsen and Stroka, 1995)
(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, 2008)
(UWGM 181) maxilla, jugal, postorbitals, squamosal, quadratojugal, quadrate, partial braincase, partial pterygoid, dentaries, splenial, surangular, prearticular, three vertebrae, 100 fragments (Larson, 2008)
(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, 2008)
(private coll.; referred to lancensis) (juvenile) teeth (with Tinker)
(private coll.; Belle) (subadult) (Blasing, DML 2006)
(private coll.) (adult) (with Tinker) maxilla, jugal (Blasing, DML 2006)
(private coll.; Ivan) about fifteen presacral vertebrae, about twenty-five presacral ribs, sacrum, abouttwenty-five caudal vertebrae, about thirty chevrons, scapulocoracoid, partial ilia, pubes, ischia, femur, tibia, fibula, astragalus, two metatarsals, six pedal phalanges (Larson, 2008)
(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, 2008)
(private coll.; Wayne) dorsal vertebra, several rib or gastralia fragments, nineteen caudal vertebrae, two chevrons, elements (Larson, 2008)
five teeth, vertebral fragment, distal metatarsal, phalanx (Lupton, Gabriel and West, 1980)
?(referred to lancensis) (juvenile) phalanx (Stenerson and O'Conner, 1994)
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; 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, 2008)
(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., 2005)
(MOR 1126; Celeste or C-rex) (~14.1 m?) surangular, prearticular, three partial dorsal vertebrae, twenty dorsal ribs, chevron (Larson, 2008)
(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, 2008)
(MOR 1131; J-rex) cranial elements including frontals, parietals, braincase (Larson, 2008)
(MOR 1152; Frank or F-rex) posterior dorsal vertebrae, posterior dorsal ribs, seven caudal vertebrae, four chevrons, pelvis, hindlimb, metatarsal (Larson, 2008)
?(juvenile and adult) ninety-one teeth (Larson, Nellermoe and Gould, 2003)
Late Maastrichtian, Late Cretaceous
Javelina Formation, Texas, US
material (Lawson, 1976; Lehman, 1985)
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, 2008)
(BMNH 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)
(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, 2008)
(MMS 51-2004) frontal, partial braincase (Molnar, 1978)
(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)
(UCMP 73081) (UCMP online)
(UCRC PV1) (~8.5 m) presacral vertebrae, dorsal ribs, gastralia, scapulocoracoids, coracoid fragments, furcula, forelimbs, hindlimb fragments (Lipkin et al., 2007)
(USNM 2110) (~12.2 m) distal metatarsal IV (~590 mm) (Gilmore, 1920)
(USNM 6183) (~9.8 m; 2.4 tons) femur (1.033 m), tibia (890 mm), proximal fibula (Gilmore, 1920)
(USNM 8064) ilium (Gilmore 1920)
?(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-PU 16516) (YPM online)
(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, 2008)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of the Kirtland Formation, New Mexico, US
(NMMNH P-7199) partial dentary, tooth fragments, partial vertebra (Carr and Williamson, 2000)
(NMMNH P-13000; = UNM FKK-076) tooth (Lucas et al., 1987)
Late Maastrichtian, Late Cretaceous
Livingston Formation, Montana, US
material (McMannis, 1965)
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 (698 mm), pedal phalanx IV-1 (53 mm), six pedal phalanges (Russell, 1970)
(uncollected) skull (Currie pers. comm. to Ford and Chure 2001)
Late Maastrichtian, Late Cretaceous
Hall Lake Member of the McRae Formation, New Mexico, US
(NMMNH P-3698; = NMMNH P-1013-1) postorbital, squamosal, palatine (missidentified as an articular), dentary, splenial, prearticular, articular, three teeth, three chevrons (Gillette, Wolberg and Hunt, 1986)
Late Maastrichtian, Late Cretaceous
North Horn Formation, Utah, US
?(UMNH 7515) ungual (Difley and Ekdale, 2002)
(UMNH 7626) partial tooth (Difley and Ekdale, 2002)
(UMNH 11000) postorbital, squamosal, third cervical vertebra, fourth cervical vertebra, dorsal rib, second sacral vertebra, third sacral vertebra, fourth sacral vertebra, six mid caudal vertebrae, six chevrons, partial ilium, proximal ischium, tibia, fibula, astragalus (Sampson and Loewen, 2005)
Late Maastrichtian, Late Cretaceous
Tornillo Formation, Texas, US
(TMM 41436-1; material of Tyrannosaurus "vannus") (subadult) maxilla (Lawson, 1976)
Late Maastrichtian, Late Cretaceous
Willow Creek Formation, Alberta, Canada
(RTMP 81.6.1; Black Beauty) (11.7 m; 3.23 tons; 18 year old adult) skull, partial mandibles (dentary 770 mm), five cervical vertebrae, two cervical ribs, seven dorsal vertebrae, eight dorsal ribs, humerus (302 mm), manual phalanx, femora (1.21 m), tibiae, fibula, astragalus, calcaneum, four metatarsals, five pedal phalanges (Currie, 1993)
Late Cretaceous
Alberta, Canada
?(referred to lancensis) skull, skeleton (Langston's 1955 field notes; www.paleofile.com)
Late Cretaceous
Saskatchewan, Canada
pedal phalanx (Langston's 1955 field notes; www.paleofile.com)
Late Cretaceous
Montana, US
(MOR 1156; J-rex2) four elements (MOR online)
(MOR 1190) phalanx (MOR online)
(MOR 1191) fibula (MOR online)
(MOR 1198; Jen-rex) femoral fragment, phalanx (MOR online)
(MOR 1602; H-rex) pedal phalanx (MOR online)
(MOR 1628) maxilla (MOR online)
?
(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. 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 BMNH 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 Osborn and Brown (1908) being due to comparisons with Euoplocephalus.
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, news reports from right before the publication of Bakker et al.'s paper erroneously 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; Larson, 2005; Witmer and Ridgely, 2005). 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 a conference held in 2005 at the museum, The Origin, Systematics and Paleobiology of Tyrannosauridae, and will be described in the future by Bakker, Larson and Currie. A portion of the BMRP's website is devoted to Jane- http://www.visitjane.com/. Evidence for CMN 7541 and BMRP 2002.4.1 being distinct from T. rex include a higher tooth count, notches in the dorsal quadratojugal, lateral pneumatic foramen on the quadratojugal and unspecified braincase morphologies. However, no Hell Creek tyrannosaurine adults with these characters are known, nor are any juveniles lacking them. I provisionally accept Nanotyrannus as a juvenile Tyrannosaurus rex, though the publication of BMRP 2002.4.1's description and papers presented at the Burpee Symposium 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 as a juvenile T. rex, but have been referred to Nanotyrannus as well (Larson pers. comm., 1997 to Ford and Chure, 2001). A lacrimal may also belong to this specimen. Around 2000, 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, 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 (nor have jugal or lacrimal differences been noted). 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.
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 has been 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 placed as 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 a 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 (2002) 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 specimen of an undescribed possibly albertosaurine tyrannosauroid) 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.
Sue or T. "stanwinstonorum"- 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 descibed, however. 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. FMNH PR2081 has a website devoted to it- http://www.fieldmuseum.org/SUE/.
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. A portion of the museum's website is devoted to the specimen- http://www.carnegiemnh.org/ditw/paleolab/samson/index.htm. Glut (2002) reported the femur is 1.36 meters long, but Larson (2008) 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 (Larson, 1994).
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 is devoted to the specimen- 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- http://www.bhigr.com/store/product.php?productid=398.
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 were present in the jackets with Tinker. Unfortunately, Tinker was not deposited in a museum and the hired preparator declared bankruptcy, so the specimen is in storage in Pennsylvania as of 2006. For a time, Harding County, SD owned Tinker, as the lease between it and the people working on the fossil was declared invalid, though the collector's regained ownership in February 2008. It's unknown if or when Tinker will be available for scientific study.
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, 2000) 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, 2000) and have a tibiofemoral ratio of 1.0. However, Larson (2008) 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) 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, 2008). The impressions appear to be scaly. As of 2004, many bones had been prepared. A website containing numerous photographs of the specimen can be seen here- 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, 2008). 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, and the Aublysodon's holotype (which Phylocode dictates the definition be based on) is indeterminate. It could be a juvenile Gorgosaurus, Daspletosaurus or even a sister taxon to T. rex. Ironically, the discovery of an apparent possible ancestor of T. rex by Stein and Triebold (2005) in the same formation as Aublysodon's holotype makes the latter situation more than hypothetical. 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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