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Rescuing orphaned refs ("FT2014" from Tyrannosauridae)
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[[File:Journal.pone.0079420.g001.tif|thumb|upright=1.5|alt=Skeletal reconstruction of two tyrannosaurs superimposed over each other, with known bones highlighted in yellow; photographs of various fossils appear below|Skeletal diagrams showing [[type (biology)|holotype]] remains of ''[[Lythronax]]'' (A) and a ''[[Teratophoneus]]'' specimen (B). N–P show selected bones of the former]]
[[File:Journal.pone.0079420.g001.tif|thumb|upright=1.5|alt=Skeletal reconstruction of two tyrannosaurs superimposed over each other, with known bones highlighted in yellow; photographs of various fossils appear below|Skeletal diagrams showing [[type (biology)|holotype]] remains of ''[[Lythronax]]'' (A) and a ''[[Teratophoneus]]'' specimen (B). N–P show selected bones of the former]]
Valid genera would not be named until the 2010s, where in 2011 announced the publication of ''[[Teratophoneus]]'' by Thomas D. Carr and colleagues. The fossils were first found in the [[Kaiparowits Formation]] of southern Utah. Later, fossils from the same formation were discovered and identified as the genus. [[Argon-argon dating|Argon-argon radiometric dating]] indicates that the Kaiparowits Formation was deposited between 76.1 and 74.0 million years ago, during the [[Campanian]] stage of the [[Late Cretaceous]] period. This date means that ''Teratophoneus'' lived in the middle of the Campanian stage of the Late Cretaceous.<ref name=":0">{{Cite journal|last1=Carr|first1=T.D.|last2=Williamson|first2=T.E.|last3=Britt|first3=B.B.|last4=Stadtman|first4=K.|date=2011|title=Evidence for high taxonomic and morphologic tyrannosaurid diversity in the Late Cretaceous (Late Campanian) of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits formation of Utah|journal=Naturwissenschaften|volume=98|issue=3|pages=241–246|doi=10.1007/s00114-011-0762-7|pmid=21253683|bibcode=2011NW.....98..241C|s2cid=13261338}}</ref> In the same year ''[[Zhuchengtyrannus]]'' was named by David W. E. Hone and colleagues based on the holotype ZCDM V0031, a nearly complete right maxilla and associated left dentary (lower jaw, both with teeth) housed at Zhucheng Dinosaur Museum.<ref name="Zhuchengtyrannus">{{cite journal|last1=Hone|first1=D. W. E.|last2=Wang|first2=K.|last3=Sullivan|first3=C.|last4=Zhao|first4=X.|last5=Chen|first5=S.|last6=Li|first6=D.|last7=Ji|first7=S.|last8=Ji|first8=Q.|last9=Xu|first9=X.|date=2011|title=A new, large tyrannosaurine theropod from the Upper Cretaceous of China|journal=Cretaceous Research|volume=32|issue=4|pages=495–503|doi=10.1016/j.cretres.2011.03.005}}</ref> In 2014 came ''[[Nanuqsaurus]]'', the northern most tyrannosaurid found in [[Prince Creek Formation]] of the [[Alaska North Slope|North Slope of Alaska]], United States.<ref name="FT2014" /><ref>{{cite journal|title=Theropod teeth from the Prince Creek Formation (Cretaceous) of northern Alaska, with speculations on Arctic Dinosaur paleoecology|date=2000|volume=20|issue=4|pages=675|doi=10.1671/0272-4634(2000)020[0675:TTFTPC]2.0.CO;2|journal=Journal of Vertebrate Paleontology|last1=Fiorillo|first1=Anthony R.|last2=Gangloff|first2=Roland A.}}</ref> In the same year also announced ''[[Qianzhousaurus]]'' known from a partial sub-adult individual consisting of a nearly complete skull with the lower jaws missing all [[teeth]] (lost during fossilization), 9 [[cervical vertebrae]], 3 [[dorsal vertebrae]], 18 caudal vertebrae, both [[scapulocoracoid]]s, partial [[ilium (bone)|ilia]], and the left hindlimb compromising the [[femur]], [[tibia]], [[fibula]], [[Talus bone|astragalus]] with [[calcaneum]], and [[metatarsals]] III and IV.<ref name=Qianzhousaurus>{{cite journal|last1=Lü|first1=J.|last2=Yi|first2=L.|last3=Brusatte|first3=S. L.|last4=Yang|first4=L.|last5=Chen|first5=L.|date=2014|title=A new clade of Asian Late Cretaceous long-snouted tyrannosaurids|journal=Nature Communications|volume=5|number=3788|bibcode=2014NatCo...5.3788L|doi=10.1038/ncomms4788|doi-access=free|pmid=24807588}}</ref> ''[[Lythronax]]'', the oldest known member of Tyrannosauridae, was described in 2013 by Mark A. Loewen and colleagues from a nearly complete specimen that was uncovered in 2009 in the [[Wahweap Formation]] of the [[Grand Staircase–Escalante National Monument]].<ref name="loewen2013">{{cite journal |last1=Loewen |first1=M.A. |last2=Irmis |first2=R.B. |last3=Sertich |first3=J.J.W. |last4=Currie |first4=P.J. |last5=Sampson |first5=S.D. |editor-last=Evans |editor-first=D.C |year=2013 |title=Tyrant dinosaur evolution tracks the rise and fall of Late Cretaceous oceans |journal=[[PLoS ONE]] |doi=10.1371/journal.pone.0079420 |volume=8 |issue=11 |pages=e79420 |ref={{sfnRef |Loewen ''et al.''|2013}} |pmid=24223179 |pmc=3819173 |bibcode=2013PLoSO...879420L|doi-access=free }}</ref>
Valid genera would not be named until the 2010s, where in 2011 announced the publication of ''[[Teratophoneus]]'' by Thomas D. Carr and colleagues. The fossils were first found in the [[Kaiparowits Formation]] of southern Utah. Later, fossils from the same formation were discovered and identified as the genus. [[Argon-argon dating|Argon-argon radiometric dating]] indicates that the Kaiparowits Formation was deposited between 76.1 and 74.0 million years ago, during the [[Campanian]] stage of the [[Late Cretaceous]] period. This date means that ''Teratophoneus'' lived in the middle of the Campanian stage of the Late Cretaceous.<ref name=":0">{{Cite journal|last1=Carr|first1=T.D.|last2=Williamson|first2=T.E.|last3=Britt|first3=B.B.|last4=Stadtman|first4=K.|date=2011|title=Evidence for high taxonomic and morphologic tyrannosaurid diversity in the Late Cretaceous (Late Campanian) of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits formation of Utah|journal=Naturwissenschaften|volume=98|issue=3|pages=241–246|doi=10.1007/s00114-011-0762-7|pmid=21253683|bibcode=2011NW.....98..241C|s2cid=13261338}}</ref> In the same year ''[[Zhuchengtyrannus]]'' was named by David W. E. Hone and colleagues based on the holotype ZCDM V0031, a nearly complete right maxilla and associated left dentary (lower jaw, both with teeth) housed at Zhucheng Dinosaur Museum.<ref name="Zhuchengtyrannus">{{cite journal|last1=Hone|first1=D. W. E.|last2=Wang|first2=K.|last3=Sullivan|first3=C.|last4=Zhao|first4=X.|last5=Chen|first5=S.|last6=Li|first6=D.|last7=Ji|first7=S.|last8=Ji|first8=Q.|last9=Xu|first9=X.|date=2011|title=A new, large tyrannosaurine theropod from the Upper Cretaceous of China|journal=Cretaceous Research|volume=32|issue=4|pages=495–503|doi=10.1016/j.cretres.2011.03.005}}</ref> In 2014 came ''[[Nanuqsaurus]]'', the northern most tyrannosaurid found in [[Prince Creek Formation]] of the [[Alaska North Slope|North Slope of Alaska]], United States.<ref name="FT2014">{{Cite journal| last1 = Fiorillo | first1 = A. R. | author-link = Anthony Fiorillo| last2 = Tykoski | first2 = R. S. | author-link2 = Ronald Tykoski| year = 2014| title = A Diminutive New Tyrannosaur from the Top of the World| editor-last = Dodson| editor-first = Peter| editor-link = Peter Dodson| journal = [[PLoS ONE]]| volume = 9| issue = 3| pages = e91287| doi = 10.1371/journal.pone.0091287| ref = {{sfnRef|Fiorillo & Tykoski|2014}} | pmid=24621577 | pmc=3951350| bibcode = 2014PLoSO...991287F| doi-access = free }}</ref><ref>{{cite journal|title=Theropod teeth from the Prince Creek Formation (Cretaceous) of northern Alaska, with speculations on Arctic Dinosaur paleoecology|date=2000|volume=20|issue=4|pages=675|doi=10.1671/0272-4634(2000)020[0675:TTFTPC]2.0.CO;2|journal=Journal of Vertebrate Paleontology|last1=Fiorillo|first1=Anthony R.|last2=Gangloff|first2=Roland A.}}</ref> In the same year also announced ''[[Qianzhousaurus]]'' known from a partial sub-adult individual consisting of a nearly complete skull with the lower jaws missing all [[teeth]] (lost during fossilization), 9 [[cervical vertebrae]], 3 [[dorsal vertebrae]], 18 caudal vertebrae, both [[scapulocoracoid]]s, partial [[ilium (bone)|ilia]], and the left hindlimb compromising the [[femur]], [[tibia]], [[fibula]], [[Talus bone|astragalus]] with [[calcaneum]], and [[metatarsals]] III and IV.<ref name=Qianzhousaurus>{{cite journal|last1=Lü|first1=J.|last2=Yi|first2=L.|last3=Brusatte|first3=S. L.|last4=Yang|first4=L.|last5=Chen|first5=L.|date=2014|title=A new clade of Asian Late Cretaceous long-snouted tyrannosaurids|journal=Nature Communications|volume=5|number=3788|bibcode=2014NatCo...5.3788L|doi=10.1038/ncomms4788|doi-access=free|pmid=24807588}}</ref> ''[[Lythronax]]'', the oldest known member of Tyrannosauridae, was described in 2013 by Mark A. Loewen and colleagues from a nearly complete specimen that was uncovered in 2009 in the [[Wahweap Formation]] of the [[Grand Staircase–Escalante National Monument]].<ref name="loewen2013">{{cite journal |last1=Loewen |first1=M.A. |last2=Irmis |first2=R.B. |last3=Sertich |first3=J.J.W. |last4=Currie |first4=P.J. |last5=Sampson |first5=S.D. |editor-last=Evans |editor-first=D.C |year=2013 |title=Tyrant dinosaur evolution tracks the rise and fall of Late Cretaceous oceans |journal=[[PLoS ONE]] |doi=10.1371/journal.pone.0079420 |volume=8 |issue=11 |pages=e79420 |ref={{sfnRef |Loewen ''et al.''|2013}} |pmid=24223179 |pmc=3819173 |bibcode=2013PLoSO...879420L|doi-access=free }}</ref>


==Systematics==
==Systematics==

Revision as of 16:55, 22 November 2021

Tyrannosaurines
Temporal range: Late Cretaceous,
80.6–66 Ma
Skeletal mount of Alioramus.
Skeletal mount of Tyrannosaurus.
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: †Pantyrannosauria
Clade: †Eutyrannosauria
Family: †Tyrannosauridae
Subfamily: †Tyrannosaurinae
Osborn, 1906
Type species
†Tyrannosaurus rex
Osborn, 1905
Subgroups[1]

The tyrannosaurines (Tyrannosaurinae) is an extinct subfamily of Tyrannosauridae a family of coelurosaurian theropod dinosaurs that consists of at least three tribes and several genera. All fossils of these genera have been found in the Late Cretaceous deposits of western North America and east Asia. Compare to the related subfamily Albertosaurinae, tyrannosaurines overall are more robust and larger though the alioramins gracile. This subfamily also includes the oldest known tyrannosaurid genus Lythronax as well as the youngest and one of the last known member of the group Tyrannosaurus.

History of discovery

Deinodon teeth, the earliest known tyrannosaurid remains

The first remains of tyrannosaurids were uncovered during expeditions led by the Geological Survey of Canada, which located numerous scattered teeth. These distinctive dinosaur teeth were given the name Deinodon ("terrible tooth") by Joseph Leidy in 1856. In 1892 Edward Drinker Cope described more tyrannosaur material in the form of isolated vertebrae, and gave this animal the name Manospondylus gigas. This discovery was mostly overlooked for over a century, and caused controversy in the early 2000s when it was discovered that this material actually belonged to, and had name priority over, Tyrannosaurus rex.[2] Later in 1905 Henry Fairfield Osborn described two tyrannosaur specimens that had been collected in Montana and Wyoming during a 1902 expedition of the American Museum of Natural History, led by Barnum Brown. Initially, Osborn considered these to be distinct species. The first, he named Dynamosaurus imperiosus ("emperor power lizard"), and the second, Tyrannosaurus rex ("king tyrant lizard"). A year later, Osborn recognized that these two specimens actually came from the same species. Despite the fact that Dynamosaurus had been found first, the name Tyrannosaurus had appeared one page earlier in his original article describing both specimens. Therefore, according to the International Code of Zoological Nomenclature (ICZN), the name Tyrannosaurus was used.[3]

The second described representative of the tyrannosaurines, Tarbosaurus (originally described as an Asiatic representative of Tyrannosaurus) was in 1955 after a large skull was recovered in a joint Soviet-Mongolian expedition to the Gobi Desert in the Mongolian Ömnögovi Province in 1946. The holotype was named as Tyrannosaurus bataar by Evgeny Maleev as Tyrannosaurus bataar.[4] The genus Tarbosaurus was also described in the same year based on PIN 551–2, a specimen with a skull and skeletal remains discovered by the same expedition in 1948 and 1949 as Tarbosaurus efremovi.[5] It was only in 1965 that Ty. bataar and Ta. efremovi were the same species, the latter being a younger animal, and distinct from Tyrannosaurus recognized by A.K. Rozhdestvensky who recombined the species as Tarbosaurus bataar.[6]

In the 1970s saw the description of two genera. In 1970 saw the publication of Daspletosaurus, with the holotype CMN 8506 consisting of a partial skeleton including the skull, the shoulder, a forelimb, the pelvis, a femur and all of the vertebrae from the neck, torso and hip, as well as the first eleven tail vertebrae. It was discovered in 1921 near Steveville, Alberta, by Charles Mortram Sternberg, who thought it was a new species of Gorgosaurus, but was found to be a new genus by Dale Russell using the aforementioned CMN 8506.[7] The second, Alioramus, described in 1976 by Sergei Kurzanov which the holotype (PIN 3141/1) is a partial skull associated with three metatarsals found by a joint Soviet-Mongolian expedition to the Gobi Desert in the early 1970s found these remains at a locality known as Nogon-Tsav in the Mongolian province of Bayankhongor, Nemegt Formation.[8]

From 1977 to 2009 saw the publications of several genera. In Asia they include Shanshanosaurus (1977)[9], Maleevosaurus (1992)[10], and Raptorex (2009)[11], while in North America saw Nanotyrannus (1988)[12], Dinotyrannus and Stygivenator (1995).[13] These genera, however are controversial as the remains of these animals are immature or juvenile individuals. As of 2021 it is widely understood that the Asian specimens are early growth stages of Tarbosaurus[14][15][16], while the North American specimens are those of Tyrannosaurus.[17][18]

Skeletal reconstruction of two tyrannosaurs superimposed over each other, with known bones highlighted in yellow; photographs of various fossils appear below
Skeletal diagrams showing holotype remains of Lythronax (A) and a Teratophoneus specimen (B). N–P show selected bones of the former

Valid genera would not be named until the 2010s, where in 2011 announced the publication of Teratophoneus by Thomas D. Carr and colleagues. The fossils were first found in the Kaiparowits Formation of southern Utah. Later, fossils from the same formation were discovered and identified as the genus. Argon-argon radiometric dating indicates that the Kaiparowits Formation was deposited between 76.1 and 74.0 million years ago, during the Campanian stage of the Late Cretaceous period. This date means that Teratophoneus lived in the middle of the Campanian stage of the Late Cretaceous.[19] In the same year Zhuchengtyrannus was named by David W. E. Hone and colleagues based on the holotype ZCDM V0031, a nearly complete right maxilla and associated left dentary (lower jaw, both with teeth) housed at Zhucheng Dinosaur Museum.[20] In 2014 came Nanuqsaurus, the northern most tyrannosaurid found in Prince Creek Formation of the North Slope of Alaska, United States.[21][22] In the same year also announced Qianzhousaurus known from a partial sub-adult individual consisting of a nearly complete skull with the lower jaws missing all teeth (lost during fossilization), 9 cervical vertebrae, 3 dorsal vertebrae, 18 caudal vertebrae, both scapulocoracoids, partial ilia, and the left hindlimb compromising the femur, tibia, fibula, astragalus with calcaneum, and metatarsals III and IV.[23] Lythronax, the oldest known member of Tyrannosauridae, was described in 2013 by Mark A. Loewen and colleagues from a nearly complete specimen that was uncovered in 2009 in the Wahweap Formation of the Grand Staircase–Escalante National Monument.[24]

Systematics

There are at least three lineages of tyrannosaurines.[25] The basal most clade is the tribe Alioramini, the second clade to diverge is an unnamed clade comprising the American southwest taxa Dynamoterror, Lythronax, and Teratophoneus. They are sister to a clade comprising of Nanuqsaurus and the tribes Daspletosaurini and Tyrannosaurini.

The cladogram below is the result of the phylogenetic analysis performed by Voris et al. (2020):[25]

Eutyrannosauria

References

  1. ^ Holtz, Thomas R. Jr. (2012) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2011 Appendix. Archived 2017-08-12 at the Wayback Machine
  2. ^ Breithaupt, B.H.; Southwell, E.H.; Matthews, N.A. (18 October 2005). "In Celebration of 100 years of Tyrannosaurus rex: Manospondylus gigas, Ornithomimus grandis, and Dynamosaurus imperiosus, the Earliest Discoveries of Tyrannosaurus rex in the West". Abstracts with Programs. 2005 Salt Lake City Annual Meeting. Vol. 37. Geological Society of America. p. 406. Archived from the original on 22 October 2019. Retrieved 8 October 2008.
  3. ^ Breithaup, BH; Southwell EH; Matthews NA (2006). "Dynamosaurus imperiosus and the earliest discoveries of Tyrannosaurus rex in Wyoming and the West". New Mexico Museum of Natural History and Science Bulletin. 35: 257–258.
  4. ^ Maleev, Evgeny A. (1955). "Giant carnivorous dinosaurs of Mongolia". Doklady Akademii Nauk SSSR. 104 (4): 634–637.
  5. ^ Maleev, E. A. (1955). "New carnivorous dinosaurs from the Upper Cretaceous of Mongolia" (PDF). Doklady Akademii Nauk SSSR. 104 (5). translated by F. J. Alcock: 779–783.
  6. ^ Rozhdestvensky, Anatoly K. (1965). "Growth changes in Asian dinosaurs and some problems of their taxonomy". Paleontological Journal. 3: 95–109.
  7. ^ Russell, Dale A. (1970). "Tyrannosaurs from the Late Cretaceous of western Canada". National Museum of Natural Sciences Publications in Paleontology. 1: 1–34.
  8. ^ Kurzanov, Sergei M. "A new carnosaur from the Late Cretaceous of Nogon-Tsav, Mongolia". The Joint Soviet-Mongolian Paleontological Expedition Transactions (in Russian). 3: 93–104.
  9. ^ Dong Zhiming (1977). "On the dinosaurian remains from Turpan, Xinjiang". Vertebrata PalAsiatica (in Chinese). 15: 59–66.
  10. ^ Carpenter, Ken. (1992). "Tyrannosaurids (Dinosauria) of Asia and North America". In Mateer, Niall J.; Peiji, Chen (eds.). Aspects of Nonmarine Cretaceous Geology. Beijing: China Ocean Press. pp. 250–268.
  11. ^ Sereno, P.; Tan, L.; Brusatte, S. L.; Kriegstein, H. J.; Zhao, X.; Cloward, K. (2009). "Tyrannosaurid skeletal design first evolved at small body size". Science. 326 (5951): 418–422. Bibcode:2009Sci...326..418S. doi:10.1126/science.1177428. PMID 19762599. S2CID 1953405.
  12. ^ Bakker, R.T.; Williams, M.; Currie, P.J. (1988). "Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana". Hunteria. 1: 1–30.
  13. ^ Olshevsky, G. (1995). "The origin and evolution of the tyrannosaurids". Kyoryugaku Saizensen [Dino Frontline]. 9–10: 92–119.
  14. ^ Carr, Thomas D. (1999). "Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)". Journal of Vertebrate Paleontology. 19 (3): 497–520. doi:10.1080/02724634.1999.10011161.
  15. ^ Currie, Philip J.; Dong Zhiming (2001). "New information on Shanshanosaurus huoyanshanensis, a juvenile tyrannosaurid (Theropoda, Dinosauria) from the Late Cretaceous of China". Canadian Journal of Earth Sciences. 38 (12): 1729–1737. Bibcode:2001CaJES..38.1729C. doi:10.1139/cjes-38-12-1729.
  16. ^ Fowler, DW; Woodward, HN; Freedman, EA; Larson, PL; Horner, JR (2011). "Reanalysis of "Raptorex kriegsteini": A Juvenile Tyrannosaurid Dinosaur from Mongolia". PLOS ONE. 6 (6): e21376. Bibcode:2011PLoSO...6E1376F. doi:10.1371/journal.pone.0021376. PMC 3126816. PMID 21738646.
  17. ^ Woodward, Holly N.; Tremaine, Katie; Williams, Scott A.; Zanno, Lindsay E.; Horner, John R.; Myhrvold, Nathan (2020). "Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy "Nanotyrannus" and supports ontogenetic niche partitioning in juvenile Tyrannosaurus". Science Advances. 6 (1): eaax6250. Bibcode:2020SciA....6.6250W. doi:10.1126/sciadv.aax6250. ISSN 2375-2548. PMC 6938697. PMID 31911944.
  18. ^ Carr, T.D. (2020). "A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence". PeerJ. 8: e9192. doi:10.7717/peerj.9192.
  19. ^ Carr, T.D.; Williamson, T.E.; Britt, B.B.; Stadtman, K. (2011). "Evidence for high taxonomic and morphologic tyrannosaurid 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. Bibcode:2011NW.....98..241C. doi:10.1007/s00114-011-0762-7. PMID 21253683. S2CID 13261338.
  20. ^ Hone, D. W. E.; Wang, K.; Sullivan, C.; Zhao, X.; Chen, S.; Li, D.; Ji, S.; Ji, Q.; Xu, X. (2011). "A new, large tyrannosaurine theropod from the Upper Cretaceous of China". Cretaceous Research. 32 (4): 495–503. doi:10.1016/j.cretres.2011.03.005.
  21. ^ Fiorillo, A. R.; Tykoski, R. S. (2014). Dodson, Peter (ed.). "A Diminutive New Tyrannosaur from the Top of the World". PLoS ONE. 9 (3): e91287. Bibcode:2014PLoSO...991287F. doi:10.1371/journal.pone.0091287. PMC 3951350. PMID 24621577.
  22. ^ Fiorillo, Anthony R.; Gangloff, Roland A. (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. doi:10.1671/0272-4634(2000)020[0675:TTFTPC]2.0.CO;2.
  23. ^ Lü, J.; Yi, L.; Brusatte, S. L.; Yang, L.; Chen, L. (2014). "A new clade of Asian Late Cretaceous long-snouted tyrannosaurids". Nature Communications. 5 (3788). Bibcode:2014NatCo...5.3788L. doi:10.1038/ncomms4788. PMID 24807588.
  24. ^ Loewen, M.A.; Irmis, R.B.; Sertich, J.J.W.; Currie, P.J.; Sampson, S.D. (2013). Evans, D.C (ed.). "Tyrant dinosaur evolution tracks the rise and fall of Late Cretaceous oceans". PLoS ONE. 8 (11): e79420. Bibcode:2013PLoSO...879420L. doi:10.1371/journal.pone.0079420. PMC 3819173. PMID 24223179.
  25. ^ a b Voris, Jared T.; Therrien, Francois; Zelenitzky, Darla K.; Brown, Caleb M. (2020). "A new tyrannosaurine (Theropoda:Tyrannosauridae) from the Campanian Foremost Formation of Alberta, Canada, provides insight into the evolution and biogeography of tyrannosaurids". Cretaceous Research. 110: 104388. doi:10.1016/j.cretres.2020.104388.
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