Abstract
Differences in proportions, anatomy, and species display features between specimens of all sizes from the latest Maastrichtian formations of the North American upper plains, greatly exceed those observed in other eutyrannosaur species, genera and subfamilies. The question is not whether more than one taxon was extant, but how many and at what ranks. Among the large Tyrannosaurus, early T. imperator retains the robust proportions, two lower functional incisors and postorbital boss features derived from earlier tyrannosaurids—the boss that includes highly atypical, large, long spindles not present in stratigraphically higher species. Among the latter, about half are robust T. rex specimens with single lower incisors, and are adorned by unique, very prominent “Mickey Mouse” postorbital display discs not previously observed in Tyrannosaurus. Contemporary T. regina skulls and skeletons are markedly more gracile, and lack either the postorbital spindles or discs. About half a dozen small remains are from juvenile specimens of Tyrannosaurus with the same tooth counts and lack of a prominent lateral dentary groove as the adults. Twice as many cannot be juveniles because their preserved hands are absolutely as large or larger than those of large adults—appendages do not atrophy with maturity in amniotes, and the feature excludes them from being tyrannosaurids. Furthermore, they have more teeth than adult specimens of Tyrannosaurus—tooth count reduction being abnormal or absent among growing reptiles. Growth arcs preserved in bone sections show these non-tyrannosaurids were growth slowing or ceasing subadults or adults, not rapidly increasing mass expansion expected in early ontogeny giants. The large-handed basal eutyrannosaurs are diverse, some but not necessarily all being Nanotyrannus or Stygivenator. Probably having evolved in Appalachia, the basal eutyrannosaurs were invading western North America (over the recently emerged land bridge), where they continued to evolve as they successfully competed with juvenile Tyrannosaurus. The result was an exceptionally diverse assemblage of tyrannosaurs at that end Cretaceous locality—caused by the rare ancient geographical event.
References
- Bakker, R.T. & Williams, M. (1988) Nanotyrannus, and new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana. Hunteria, 1, 1–30. https://doi.org/10.1075/z.31.02int
- Botho, J., Choiniere, J.N. & Benson, R.B.J. (2022) Rapid growth preceded gigantism in sauropodomorph evolution. Current Biology, 32, 4501–4507. https://doi.org/10.1016/j.cub.2022.08.031
- Brochu, C.A. (2003) Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology, 22 (7), 1–138. https://doi.org/10.1080/02724634.2003.10010947
- Brown, C.M., Currie, P.J. & Therrien, F. (2022) Intraspecific facial bite marks in tyrannosaurids provide insight into sexual maturity and evolution of bird-like intersexual display. Paleobiology, 48, 2–43. https://doi.org/10.1017/pab.2021.29
- Brown, C.M., VanBuren, C.S., Larson, D.W., Brink, K.S., Campione, N.E., Vavrek, M.J. & Evans, D.C. (2015) Tooth counts through growth in diapsid reptiles: implications for interpreting individual and size-related variation in the fossil record. Journal of Anatomy, 226, 322–333. https://doi.org/10.1111/joa.12280
- Brusatte, S.L. & Carr, T.D. (2016) The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports, 6, 20252. https://doi.org/10.1038/srep20252
- Brusatte, S.L., Benson, R.B.J. & Norell, M.A. (2011) The anatomy of Dryptosaurus aquilunguis (Dinosauria: Theropoda) and a review of its tyrannosauroid affinities. American Museum Novitates, 3717, 1–53. https://doi.org/10.1206/3717.2
- Brusatte, S.L., Carr, T.D. & Norell, M.A. (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, 1–197. https://doi.org/10.1206/770.1
- Brusatte, S.L., Carr, T.D., Erickson, G.M., Norell, M.A. (2009) A long-snouted, multihorned tyrannosaurid from the Late Cretaceous of Mongolia. Proceedings of the National Academy of Sciences, 106, 17261–17266. https://doi.org/10.1073/pnas.0906911106
- Brusatte, S.L., Carr, T.D., Erickson, G.M., Bever, G.S. & Norell, M.A. (2010) Tyrannosaur paleobiology: new research on ancient exemplar organisms. Science, 329, 1481–1485. https://doi.org/10.1073/pnas.0906911106
- Brusatte, S.L., Carr, T.D., Williamson, T.E., Holtz, T.R., Hone, W.E. & Williams, S.A. (2016) Dentary groove morphology does not distinguish “Nanotyrannus” as a valid taxon of tyrannosauroid dinosaur. Comment on: “Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker etal., 1988”. Cretaceous Research, 65, 232–237. https://doi.org/10.1016/j.cretres.2016.02.007
- Burnham, D.A., Atkins-Weltman, K.L. & Jevnikar, E.M. (2018) A new juvenile Tyrannosaurus rex from the Hell Creek Formation of eastern Montana provides insights into cranial and dental ontogeny. Society of Vertebrate Paleontology 78th Annual Meeting, 99.
- Canale, J.I., Apesteguia, S., Gallina, P.A., Mitchell, J., Smith, N.D., Cullen, T.M., Shinya, A., Haluza, A., Gianechini, F.A. & Makovicky, P.J. (2022) New giant carnivorous dinosaur reveals convergent evolutionary trends in theropod evolution. Current Biology, 32, 3195–3202.e5. https://doi.org/10.1016/j.cub.2022.05.057
- Carpenter, K., Russell, D.A., Baird, D. & Denton, R. (1997) Redescription of the holotype of Dryptosaurus aquilunguis from the Upper Creteacous of New Jersey. Journal of Vertrebrate Paleontology, 17, 561–573. https://doi.org/10.1080/02724634.1997.10011003
- Carr, T.D. (1999) Craniofacial ontogeny in tyrannosaurid dinosaurs. Journal of Vertebrate Paleontology, 19, 497–523. https://doi.org/10.1080/02724634.1999.10011161
- Carr, T.D. (2020) A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ, 8, e9192. https://doi.org/10.7717/peerj.9192
- Carr, T.D. & Williamson, T.E. (2010) Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosaurid from New Mexico and the origin of deep snouts in Tyrannosauroidea. Journal of Vertebrate Paleontology, 30, 1–16. https://doi.org/10.1080/02724630903413032
- Carr, T.D., Williamson, T.E., Britt, B.B. & Stadtman, K. (2011) Evidence for high taxonomic and morphologic tyrannosaurid diversity in the Late Cretaceous of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits formation of Utah. Naturwissenschaften, 98, 241–246. https://doi.org/10.1007/s00114-011-0762-7
- Carr, T.D., Varricchio, D.J., Sedlmayr, J.C., Roberts, E.M. & Moore, J.R. (2017) A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports, 7, 44942. https://doi.org/10.1038/srep44942
- Carr, T.D., Napoli, J.G., Brusatte, S.L., Holtz, T.R., Hone, D.W., Williamson, T.E. & Zanno, L.E. (2022) Insufficient evidence for multiple species of Tyrannosaurus in the latest Cretaceous of North America: A comment on “The tyrant lizard king, queen and emperor: Multiple lines of evidence support subtle evolution and probable speciation within the North American genus Tyrannosaurus”. Evolutionary Biology, 49, 327–341. https://doi.org/10.1007/s11692-022-09573-1
- Chure, D.J. & Loewen, M.A. (2020) Cranial anatomy of Allosaurus jimmadseni, a new species from the lower Morrison Formation of western North America. PeerJ, 8, e7803. https://doi.org/10.7717/peerj.7803
- Colbert, E.H. 1989. The Triassic dinosaur Coelophysis. Museum of Northern Arizona Bulletin, 57, 1–160.
- Condamine, F.L., Guinot, G., Benton, M.J. & Currie, P.J. (2021) Dinosaur biodiversity declined well before the asteroid impact, influenced by ecological and environmental pressures. Nature Communication, 12, 3833. https://doi.org/10.1038/s41467-021-23754-0
- Cullen, T.M., Canale, J.I., Apesteguia, S., Smith, N.D., Hu, D. & Makovicky, P.J. (2020) Osteohistological analyses reveal diverse strategies of theropod body-size evolution. Proceedings of the Royal Society, 287, 2020–2258. https://doi.org/10.1098/rspb.2020.2258
- Currie, P.J. (2003a) Cranial anatomy of tyrannosaurids from the Late Cretaceous of Alberta. Acta Palaeontologica Polonica, 48, 191–226.
- Currie, P.J. (2003b) Allometric growth in tyrannosaurids from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences, 40, 651–665. https://doi.org/10.1139/e02-083
- Dalman, S.G., Loewen, M.A., Pyron, R.A., Jasinski, S.E., Malinzak, D.E., Lucas, S.G., Fiorillo, A.R., Currie, P.J. & Longrich, N.R. (2024) A giant tyrannosaur from the Campanian-Maastrichtian of southern North America and the evolution of tyrannosaurid gigantism. Scientific Reports, 13, 22124. https://doi.org/10.1038/s41598-023-47011-0
- Danison, A., Wedel, M., Barta, D., Woodward, H., Flora, H., Lee, A. & Snively, E. (2024) Chimerism in specimens referred to Saurophaganax maximus reveals of new species of Allosaurus. Vertebrate Anatomy Morphology Palaeontology, 12, 81–114. https://doi.org/10.18435/vamp29404
- Dow, R.A., Choong, c.y., grinang, j., lupiyaningdyah, p., ngiam, r.w.j. & kalkman, v.j. (2024) Checklist of the Odonate (Insecta) of Sundaland and Wallacea (Malaysia, Singapore, Brunei, Indonesia and Timor Leste). Zootaxa, 5460 (1), 1–122. https://doi.org/10.11646/zootaxa.5460.1.1
- Eberth, D.A. & Kamo, S.L. (2019) First high-precision U-Pb CA-ID-TIMS age for the Battle Formation, Red Deer River Valley, Alberta, Canada: Implications for ages, correlations, and dinosaur biostratigraphy of the Scollard, Frenchman, and Hell Creek Formations. Canadian Journal of Earth Sciences, 56, 1041–1051. https://doi.org/10.1139/cjes-2018-0098
- Engelman, R.K. (2024) Reconstructing Dunkleosteus terrelli (Placodermi: Arthrodira): A new look for an iconic Devonian predator. Palaeontologica Electronica, 27.3.a45. https://doi.org/10.26879/1343
- Erickson, G.M., Makovicky, P.J., Currie, P.J., Norell, M.A., Yerby, S.A. & Brochu, C.A. (2004) Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature, 430, 772–775. https://doi.org/10.1038/nature16487
- Ezcurra, M.D. & Butler, R.J. (2015) Post-hatchling cranial ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy, 226, 387–402. https://doi.org/10.1111/joa.12300
- Fowler, D.W. (2017) Revised geochronology, correlation, and dinosaur stratigraphic ranges, correlation, and dinosaur stratigraphic ranges of the Santonian–Maastrichtian formations of the Western Interior of North America. PLoS One, 12, e0188426. https://doi.org/10.1371/journal.pone.0188426
- Fowler, D.W., Woodward, H.N., Freedman, E.A., Larson, P.L. & Horner, J.R. (2011) Reanalysis of “Raptorex kriegsteini”: a juvenile dinosaur from Mongolia. PLoS One, 6, e21376. https://doi.org/10.1371/journal.pone.0021376
- Gates, T.A., Prieto-Marquez, A. & Zanno, L.E. (2012) Mountain building triggered Late Cretaceous megaherbivore dinosaur radiation. PLoS One, 7, e42135. https://doi.org/10.1371/journal.pone.0042135
- Gignac, P. & Erickson, G.M. (2017) The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports, 7, 2012. https://doi.org/10.1038/s41598-017-02161-w
- Gillette, D.D., Wolberg, D.L. & Hunt, A.P. (1986) Tyrannosaurus rex from the McRae Formation, Elephant Butte Reservoir, Sierra County, New Mexico. 235–238. In: Clemons, R.E., King, W.E., Mack, G.H. & Zidek, J. (Eds), New Mexico Geological Society 37th Annual Fall Field Conference Guidebook. Truth or Consequences Region, New Mexico Geological Society, Socorro. https://doi.org/10.56577/FFC-37.235
- Gilmore, C.W. (1946) A new carnivorous dinosaur from the Lance formation of Montana. Smithsonian Miscellaneous Collections, 106 (13), 1–19.
- Gourvennec, R. (2012) Inflation in the number of monospecific genera in brachiopods: progress or regression? A trivial approach to a real problem. Carnets de Geologie, 285–292. https://doi.org/10.4267/2042/48396
- Green, T.L., Kay, D.I. & Gignac, P.M. (2022) Intraspecific variation and directional casque asymmetry in adult southern cassowaries (Casuarius casuarius). Journal of Anatomy, 241, 951–965. https://doi.org/10.1111/joa.13733
- Griffin, C., Poust, A.W., Bugos, J.E, Morris, Z.S., Petermann, H., Fabbri, M. & Colleary, C. (2024) Assessing the ontogenetic maturity of the ‘Nanotyrannus lancensis’ holotype with hyoid osteohistology. The Society of Vertebrate Paleontology 84th Meeting Abstracts, 232–233.
- Guinard, G. (2020) Forelimb shortening of Carcharodontosauria (Dinosauria: Theropoda): an update on evolutionary anterior micromelias in non-avian theropods. Zoology, 139, 125756. https://doi.org/10.1016/j.zool.2020.125756
- Happ, J. (2008) An analysis of predator-prey behavior in a head-to-head encounter between Tyrannosaurus rex and Triceratops. In: Larson, P. & Carpenter, K. (Eds), Tyrannosaurus rex: The Tyrant King. Indiana University Press, 355–368.
- Harvati, K. & Ackermann, R.R. (2022) Merging morphological and genetic evidence to assess hybridization in western Eurasian late Pleistocene hominins. Nature Ecology & Evolution, 6, 1573–1585. https://doi.org/10.1038/s41559-022-01875-z
- Henderson, M.D. & Harrison, H. (2008) Taphonomy and environment of deposition of juvenile tyrannosaurid skeleton from the Hell Creek Formation of southeastern Montana. In: Larson, P. & Carpenter, K. (Eds), Tyrannosaurus rex: The Tyrant King. Indiana University Press, pp. 82–90.
- Horner, J.R. & Padian, K. (2004) Age and growth dynamics of Tyrannosaurus rex. Proceedings of the Royal Society B, 271, 1875–1880. https://doi.org/10.1098/rspb.2004.2829
- Hurum, J.H. & Sabath, K. (2003) Giant theropod dinosaurs from Asia and North America: Skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Paleontologica Polonica, 48, 161–190.
- Hutchinson, J.R., Bates, K.T., Molnar, J., Allen, V. & Makovicky, P.J. (2011) A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLoS One, 6, e26037. https://doi.org/10.1371/journal.pone.0026037
- Jevnikar, E.M. & Zanno, L.E. (2021) Bimodal trajectories and unresolved early growth stages in Tyrannosaurus rex growth. Society of Vertebrate Paleontology 81st Annual Meeting, 150–151.
- Knutsen, E.M. (2012) A taxonomic revision of the genus Pliosaurus. Norwegian Journal of Geology, 92, 259–276.
- Larramendi, A., Paul, G.S. & Hsu, S.Y. (2021) A review and reappraisal of the specific gravities of present and past multicellular organisms, with an emphasis on tetrapods. The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology, 304, 1833–1888. https://doi.org/10.1002/ar.24574
- Larson, N. (2008) One hundred years of Tyrannosaurus rex: The skeletons. In: Larson, P. & Carpenter, K. (Eds), Tyrannosaurus rex: The Tyrant King. Indiana University Press, Bloomington, pp. 1–55. https://doi.org/10.2979/5139.0
- Larson, P. (2008) Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson, P. & Carpenter, K. (Eds), Tyrannosaurus rex: The Tyrant King. Indiana University Press, Bloomington, pp. 103–130. https://doi.org/10.2979/5139.0
- Larson, P. (2013a) The validity of Nanotyrannus lancensis. Society of Vertebrate Paleontology 73rd Annual Meeting, 159.
- Larson, P. (2013b) The case for Nanotyrannus. In: Parrish, J.M., Molnar, R.E., Currie, P.J. & Koppelhus, E.B. (Eds), Tyrannosaurid Paleobiology. Indiana University Press, Bloomington, 15–53.
- Liberona, J.P., Soto-Acuria, S., Mendez, M.A. & Vargas, A.O. (2019) Assessment and interpretations of negative forelimb allometry in the evolution of non-avian Theropoda. Frontiers in Zoology, 16, 44. https://doi.org/10.1186/s12983-019-0342-9
- Lipkin, C., Sereno, P.C. & Horner, J.R. (2007) The furcula in Suchomimus tenerensis and Tyrannosaurs rex (Dinosauria: Theropoda: Tetanurae). Journal of Paleontology, 81, 1523–1525. https://doi.org/10.1666/06-024.1
- Longrich, N.R. & Saitta, E.T. (2024) Taxonomic status of Nanotyrannus lancensis (Dinosauria: Tyrannosauroidea)—a distinct taxon of small-bodied tyrannosaur. Fossil Studies, 2, 1–25. https://doi.org/10.3390/fossils2010001
- Lü, J.C., Yi, L., Brusatte, S.L., Yang, L. Li, H. & Chen, L. (2014) A new clade of Asian Late Cretaceous long-snouted tyrannosaurids. Nature Communications, 5, 3788. https://doi.org/10.1038/ncomms4788
- Macdonald, I. & Currie, P.J. (2018) Description of a partial Dromiceiomimus (Dinosauria: Theropoda) skeleton with comments on the validity of the genus. Canadian Journal of Earth Sciences, 56, 129–157. https://doi.org/10.1139/cjes-2018-0162
- Madsen, J.H. (1976) Allosaurus fragilis: A revised osteology. Utah Geological and Mineral Survey Bulletin, 109, 1–163.
- Maisch, M.W. (2008) Revision der Gattung Stenopterygius Jaekel, 1904 emend. VON HUENE, 1922 (Reptilia: Ichthyosauria) aus dem unteren Jura Westeuropas. Palaeodiversity, 1, 227–271.
- Maleev, E.A. (1955) Giant carnivorous dinosaurs of Mongolia. Doklady, Academy of Sciences USSR, 104, 634–637.
- Maleev, E.A. (1974) Giant carnosaurs of the family Tyrannosauridae. Joint Soviet-Mongolian Paleontological Expedition, 1, 132–191.
- Mallon, J.C. (2019) Competition structured a Late Cretaceous megaherbivorous dinosaur assemblage. Science Reports, 9, 15447. https://doi.org/10.1038/s41598-019-51709-5
- Mallon, J.C., Roloson, M., Bamforth, E., Scannella, J.B. & Ryan, M.J. (2025) The Canadian fossil record supports anagenesis in Triceratops (Ornithischia, Ceratopsia). Canadian Journal of Earth Sciences. https://doi.org/10.1139/cjes-2024-0170
- Maxwell, E.E. (2012) New metrics to differentiate species of Stenopterygius (Reptilia: Ichthyosauria) from the Lower Jurassic of southwestern Germany. Journal of Paleontology, 86, 105–115. https://doi.org/10.1666/11-038.1
- Molnar, R.E. (1978) A new theropod dinosaur from the Upper Cretaceous of central Montana. Journal of Paleontology, 52, 73–82.
- Molnar, R.E. (1980) An albertosaur from the Hell-Creek formation of Montana. Journal of Paleontology, 54, 102–108.
- Molnar, R.E. (1991) The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A, 217, 137–176.
- Myhrvold, N.P. (2013) Revisiting the estimation of dinosaur growth rates. PLoS One, 8, e81917. https://doi.org/10.1371/journal.pone.0081917
- Napoli, J.G., Powers, M.J., Ruebenstahl, A.A., Voris, J.T., Green, T.L., Turner, A.H., Flynn, J.J., Currie, P.J. & Norell, M.A. (2023) Cryptic species in the theropod fossil record. Society of Vertebrate Paleontology 83rd Annual Meeting, 319.
- Nowak, R.M. (1991) Walker’s mammals of the world V 2. Johns Hopkins University Press, 1–1629.
- O’Connor, R.E., Romanov, M.N., Kiazim, L.G., Barrett, P.M., Farré, M., Damas, J., Ferguson-Smith, M., Valenzuela, N., Larkin, D.M. & Griffin, D. (2018) Reconstruction of the diapsid ancestral genome permits chromosome evolution tracing in avian and non-avian dinosaurs. Nature Communications, 9, 1883. https://doi.org/10.1038/s41467-018-04267-9
- Olshevsky, G. & Ford, T.L. (1995) The origin and evolution of the Tyrannosauridae, part 2. Dino Frontline, 6, 75–99.
- Osborn, H.F. (1905) Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bulletin of the American Museum of Natural History, 21, 259–265.
- Osborn, H.F. (1906) Tyrannosaurus, Upper Cretaceous carnivorous dinosaur (Second Communication). Bulletin of the American Museum of Natural History, 22, 281–296.
- Osborn, H.F. (1912) Crania of Tyrannosaurus and Allosaurus. Memoirs of the American Museum of Natural History, 1, 1–30.
- Osborn, H.F. (1916) Skeletal adaptations of Ornitholestes, Struthiomimus, Tyrannosaurus. Bulletin of the American Museum of Natural History, 35, 733–771.
- Paterna, A. & Cau, A. (2022) New giant theropod material from the Kem Kem compound assemblage with implications on the diversity of the mid-Cretaceous carcharodontosaurids from North Africa. Historical Biology, 35, 2036–2044. https://doi.org/10.1080/08912963.2022.2131406
- Paul, G.S. (1988) Predatory Dinosaurs of the World. Simon & Schuster, 110–113.
- Paul, G.S. (1997) Dinosaur models: the good, the bad, and using them to estimate the mass of dinosaurs. In: Wolberg, D.L., Stump, E. & Rosenberg, G.D. (Eds), Dinofest. International Symposium Proceedings, Academy of Natural Sciences, Philadelphia. Philadelphia, Academy Sciences of Natural Sciences, pp. 129–154.
- Paul, G.S. (2008) The extreme lifestyles and habits of the gigantic tyrannosaurid superpredators of the Late Cretaceous of North America and Asia. In: Larson, P. & Carpenter, K. (Eds), Tyrannosaurus rex: The Tyrant King. Indiana University Press, Bloomington, pp. 307–352.
- Paul, G.S. (2010) The Princeton field guide to dinosaurs. Princeton University Press, 1–320. https://doi.org/10.1515/9781400836154
- Paul, G.S. (2016) The Princeton field guide to dinosaurs (Second Edition). Princeton University Press, 1–360. https://doi.org/10.1515/9781400883141
- Paul, G.S. (2018) Nonornithischian dinosaurs did to have lips, probably big lips, here’s why. Prehistoric Times, 127, 44–49.
- Paul, G.S. (2019) Determing the largest known land animal: A critical comparison of differing methods for restoring the volume and mass of extinct animals. Annals of Carnegie Museum, 85, 335–358. https://doi.org/10.2992/007.085.0403
- Paul, G.S. (2022) The Princeton field guide to pterosaurs. Princeton University Press, 1–184. https://doi.org/10.1515/9780691232218
- Paul, G.S. (2024a) The Princeton field guide to dinosaurs 3rd ed. Princeton University Press, 1–384. https://doi.org/10.1515/9780691253336
- Paul, G.S. (2024b) The Princeton field guide to predatory dinosaurs. Princeton University Press, 1–256. https://doi.org/10.1515/9780691253336
- Paul, G.S., Persons, W.S. & Van Raalte, J. (2022) The tyrant lizard king, queen and emperor: Multiple lines of evidence support subtle evolution and probable speciation within the North American genus Tyrannosaurus. Evolutionary Biology, 49, 156–179. https://doi.org/10.1007/s11692-022-09561-5
- Paulina-Carabajal, A.P., Currie, P.J., Dudgeon, T.W., Larsson, H.C.E. & Miyashita, T. (2021) Two braincases of Daspletosaurus (Theropoda: Tyrannosauridae): anatomy and comparison. Canadian Journal of Earth Sciences, 58, 885–910. https://doi.org/10.1139/cjes-2020-0185
- Persons, W.S., Currie, P.J. & Erickson, G.M. (2019) An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record, 303, 656–672. https://doi.org/10.1002/ar.24118
- Rozhdestvensky, A.K. (1965) Growth changes in Asian dinosaurs and some problems of their taxonomy. Paleontologicheskii Zhurnal, 1965, 95–109.
- Rinehart, L.F., Lucas, S.G., Heckert, A.B., Spielmann, J.A. & Celeskey, M.D. (2009) The paleobiology of Coelophysis bauri from the Upper Triassic Whitaker quarry, New Mexico, with a detailed analysis of a single quarry block. The New Mexico Museum of Natural History & Science, 45, 1–260.
- Russell, D.A. (1970) Tyrannosaurs from the Late Cretaceous of western Canada. National Museum of Natural Sciences Publications in Paleontology, 1, 1–34.
- Ryan, M.J. & Evans, D.C. (2005) Ornithischian Dinosaurs. In: Currie, P.J. & Koppelhus, E.B (Eds). Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed. Indiana University Press, pp. 312–348).
- Sampson, S.D. & Loewen, M.A. (2005) Tyrannosaurus rex from the upper Cretaceous North Horn Formation of Utah: Biostratigraphic and paleoecological implications. Journal of Vertebrate Paleontology, 25, 469–472. https://doi.org/10.1671/0272-4634(2005)025[0469:TRFTUC]2.0.CO;2
- Scannella, J.B, Fowler, D.W., Goodwin, M.B. & Horner, J.R. (2014) Evolutionary trends in Triceratops from the Hell Creek Formation, Montana. PNAS, 111, 10245–10250. https://doi.org/10.1073/pnas.1313334111
- Scherer, C.R. (2025) Multiple lines of evidence support anagenesis in Daspletosaurus and cladogenesis in derived tyrannosaurines. Cretaceous Research, 155, 106080. https://doi.org/10.1016/j.cretres.2025.106080
- Scherer, C.R. & Voiculescu-Holvad, C. (2024) Re-analysis of a dataset refutes claims of anagenesis within Tyrannosaurus-line tyrannosaurines (Theropoda, Tyrannosauridae). Cretaceous Research, 155, 105780. https://doi.org/10.1016/j.cretres.2023.105780
- Schmerge, J.D. & Rothschild, B.R. (2016a) Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus Nanotyrannus. Cretaceous Research, 61, 26–33. https://doi.org/10.1016/j.cretres.2015.12.016
- Schmerge, J.D. & Rothschild, B.R. (2016b) When a groove is not a groove: Clarification of the appearance of the dentary groove in tyrannosauroid theropods and the distinction between Nanotyrannus and Tyrannosaurus. Reply to Comment on: “Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al., 1988”. Cretaceous Research, 65, 238–243. https://doi.org/10.1016/j.cretres.2016.04.015
- Schweitzer, M.H., Zheng, W., Zanno, L., Werning, S. & Sugiyama, T. (2016) Chemistry supports the identification of gender-specific reproductive tissue in Tyrannosaurus rex. Scientific Reports, 6, 23099.
- Son, M., Erickson, G.M., Zhou, C.F., Yin, Y.L. & Makovicky, P. (2024) Intra- and inter- specific variation in Psittacosaurus. The Society of Vertebrate Paleontology 84th Meeting Abstracts, 512–513.
- Stein, W.W. & Triebold, M. (2013) Preliminary analysis of a sub-adult tyrannosaurid skeleton from the Judith River Formation of Petroleum County, Montana. In: Parrish, J.M., Molnar, R.E., Currie, P.J. & Koppelhus, E.B. (Eds), Tyrannosaurid paleobiology. Indiana University Press, Bloomington, pp. 55–77.
- Stock, J.C., Powers, M.J., Coppock, c. Raun, G.S., Sharpe, H.S., Henry, S., Jared, T., Napoli, J.G. & Currie, P.J. (2024) New species of Albertosaurus from a distinct stratigraphic unit of the Horseshoe Canyon Formation corresponding to faunal and climatic changes. The Society of Vertebrate Paleontology 84th Meeting Abstracts, 518–519.
- Therrien, F., Zelenitsky, D.K., Tanaka, K., Voris, J.T., Erickson, G.M., Currie, P.J., Debuhr, C.L. & Kobayashi, Y. (2023) Exceptionally preserved stomach contents of a young tyrannosaurid reveal an ontogenetic dietary shift in an iconic extinct predator. Science Advances, 9, eadi0505. https://doi.org/10.1126/sciadv.adi0505
- Tsuihiji, T., Watabe, M., Tsogtbaatar, K., Tsubamoto, T., Barscold, R., Suzuki, S., Lee, A.H., Ridgely, R.C., Kawahara, Y. & Witmer, L.M. (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, 497–517. https://doi.org/10.1080/02724634.2011.557116
- Wang, X., Stiegler, J., Amiot, R., Wang, X., Du, G.H., Clark, J.M. & Xu, X. (2017) Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology, 27, 144–148. https://doi.org/10.1016/j.cub.2016.10.043
- Warshaw, E.A. & Fowler, D.W. (2022) A transitional species of Daspletosaurus Russell, 1970 from the Judith River Formation of eastern Montana. PeerJ, 10, e14461. https://doi.org/10.7717/peerj.14461
- Warshaw, E.A., Barrera, G.D. & Fowler, D.W. (2024) Anagenesis and the tyrant pedigree: a response to “Re-analysis of a dataset refutes claims of anagenesis within Tyrannosaurus-line tyrannosaurines”. Cretaceous Research, 155, 105957. https://doi.org/10.1016/j.cretres.2024.105957
- Wick, S.L. (2014) New evidence for the possible occurrence of Tyrannosaurus in west Texas, and discussion of Maastrichtian tyrannosaurid dinosaurs from Big Bend National Park. Cretaceous Research, 50, 52–58. https://doi.org/10.1016/j.cretres.2014.03.010
- Witmer, L.M. & Ridgely, R.C. (2010) The Cleveland tyrannosaur skull (Nanotyrannus or Tyrannosaurus): New findings based on CT scanning, with special reference to the braincase. Kirtlandia, 57, 61–81.
- Witton, M.P. (2022) Tyannouroboros: how everything old is new again in recent proposals of Tyrannosaurus taxonomy. Mark P. Witton’s blog, http://markwitton-com.blogspot.com/2022/03/tyrannouroboros-how-everything-old-is.html.
- Woodward, H.N., Tremaine, K., Williamson S.A., Zanno, L.E., Horner, J.R. & Myhrvold, N. (2020) Growing up Tyrannosaurus rex: Osteohistory refutes the pygmy “Nanotyrannus” and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances, 6, eaax6250. https://doi.org/10.1126/sciadv.aax6250