Tyrannosaurinae

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Tyrannosaurinae
Temporal range: Late Cretaceous,
81.9–66  Ma
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(possible Cenomanian occurrence of Labocania )
Alioramus remotus Skull.jpg
Skull of Alioramus .
Tyrannosaurus rex theropod dinosaur (Hell Creek Formation, Upper Cretaceous; near Faith, northwestern South Dakota, USA) 2 (15157270200).jpg
Skull of Tyrannosaurus .
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family:† Tyrannosauridae
Subfamily:† Tyrannosaurinae
Osborn, 1906
Type genus
† Tyrannosaurus
Osborn, 1905
Subgroups [1]

Tyrannosaurinae (or tyrannosaurines) is one of the two extinct subfamilies of Tyrannosauridae, a family of coelurosaurian theropods 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. Compared to the related subfamily Albertosaurinae, tyrannosaurines overall are more robust and larger though the alioramins were gracile by comparison. This subfamily also includes Lythronax , one of the oldest known tyrannosaurid genera, as well as the youngest and most famous member of the group, Tyrannosaurus rex .

Contents

History of discovery

Deinodon teeth, the earliest known tyrannosaurid remains Deinodon.JPG
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 . [5] 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. [6]

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. [7] 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. [8] 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. [9]

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. [10] 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. [11]

From 1977 to 2009 saw the publications of several genera. In Asia they include Shanshanosaurus (1977), [12] Maleevosaurus (1992), [13] and Raptorex (2009), [14] while in North America saw Nanotyrannus (1988), [15] Dinotyrannus and Stygivenator (1995). [16] These genera, however are controversial as the remains of these animals are immature or juvenile individuals. With the possible exception of Raptorex, [17] it is widely assumed that the Asian specimens are early growth stages of Tarbosaurus, [18] [19] [20] whereas the North American specimens are those of Tyrannosaurus. [21] [22]

Skeletal diagrams showing holotype remains of Lythronax (A) and a Teratophoneus specimen (B). N-P show selected bones of the former Journal.pone.0079420.g001.tif
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. [23] 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. [24] In 2014 came Nanuqsaurus , the northern most tyrannosaurid found in Prince Creek Formation of the North Slope of Alaska, United States. [25] [26] 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. [27] 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. [28]

Natural history

Description

Size of A. remotus compared with a human Alioramus skeletal steveoc.png
Size of A. remotus compared with a human
The specimens "Sue", AMNH 5027, "Stan", and "Jane", to scale with a human. Tyrannosaurus specimens.svg
The specimens "Sue", AMNH 5027, "Stan", and "Jane", to scale with a human.

In comparison to the albertosaurines, tyrannosaurines were more heavily built and larger. The alioramin genera of Qianzhousaurus and Alioramus, however, were the exception, as they were more comparable in built to albertosaurines and have longirostrine snouts. [27] Like albertosaurines, tyrannosaurines also had heterodont dentition, large heads design to catch and kill their prey, and short didactyl arms. Based on the growth stages of Tyrannosaurus (and possibly Tarbosaurus [20] ), tyrannosaurines undergone ontogenetic changes from gracile or slender, semi-longirostrine immatures to robust, heavy-headed adults. This implies that these animals occupy different ecological niches as they developed. [22] While there is fossil evidence of earlier tyrannosauroids having feathers, [29] [30] the evidence of such structures in tyrannosaurids is controversial as a study in 2017 from Bell and colleagues found no support in feathered integument in tyrannosaurids. [31] The study used skin impressions which are small, found widely dispersed across the post-cranium at different regions of the body with a pattern similar to crocodiles. Further the croc analogy Thomas Carr and colleagues in 2017 by studying the snout of Daspletosaurus suggested they have large scales with sensory sensory neuron pits under the skin. [32] This notion has been challenged from other authors who suggested a more lip-covering of the teeth. [33]

Distribution

The temporal range for tyrannosaurines went from almost 80.6 million years ago in the Campanian stage of the Late Cretaceous epoch to 66 million years ago in the Maastrichtian stage. Fossils have been found in different formations in what is now east Asia and western North America. While the Asian alioramins are the basal most group of the tyrannosaurines, the geographic placement of albertosaurines and other eutyrannosaurian tyrannosauroids found in North America suggests greatly the tyrannosaurines are North American in origin. [34]

Systematics

Prior to the 2010s, the relationships of the tyrannosaurines was best understood as Tyrannosaurus being a sister taxon to Tarbosaurus. These two genera in turn were the sister taxon to Daspletosaurus, follow by Alioramus. There was an alternative hypothesis from Phil Currie and colleagues (2003) who suggested Daspletosaurus to be more closely related to Tarbosaurus and Alioramus than to Tyrannosaurus based on cranial features. [35] This relationship, however, has not been found in more recent studies. [23] [25] [28] [34] [36] In 1988 Gregory Paul considered all tyrannosaurines at the time except Alioramus to be species of Tyrannosaurus. [37] In the second edition of The Princeton Field Guide to Dinosaurs published in 2016, Paul would continue this thought as well as including Bistahieversor , Teratophoneus, Lythronax, and Nanuqsaurus into the genus as well. [38] This multispecies Tyrannosaurus classification is, however, not widely accepted by most paleontologists. [36] In some phylogenetic studies Bistahieversor is nested within Tyrannosaurinae, [25] [28] but it is most often recovered as the sister taxon to Tyrannosauridae instead. [34] [36]

The cladogram below displays the position of the Tyrannosaurinae within Eutyrannosauria and Tyrannosauridae, based on the results of phylogenetic analyses performed by Voris et al. (2020): [36]

Eutyrannosauria

As of 2023, at least three lineages of tyrannosaurines have been suggested. [36] The basalmost clade is the Alioramini. The second clade to diverge is the Teratophoneini, which comprises the American southwest taxa Dynamoterror, Lythronax, and Teratophoneus. They are sister to a third clade comprising Nanuqsaurus and the clade containing Daspletosaurini, which includes Daspletosaurus and Thanatotheristes, and the Tyrannosaurini, which includes Zhuchengtyrannus, Tarbosaurus, and Tyrannosaurus. [3]

The cladogram below displays the results of the strict consensus phylogenetic analysis performed by Scherer & Voiculescu-Holvad (2023), indicating the distinct lineages of tyrannosaurines. [3]

Tyrannosaurinae

Related Research Articles

<i>Tyrannosaurus</i> Genus of Late Cretaceous theropod

Tyrannosaurus is a genus of large theropod dinosaur. The type species Tyrannosaurus rex, often shortened to T. rex or colloquially T-Rex, is one of the best represented theropods. It lived throughout what is now western North America, on what was then an island continent known as Laramidia. Tyrannosaurus had a much wider range than other tyrannosaurids. Fossils are found in a variety of rock formations dating to the latest Campanian-Maastrichtian ages of the late Cretaceous period, 72.7 to 66 million years ago. It was the last known member of the tyrannosaurids and among the last non-avian dinosaurs to exist before the Cretaceous–Paleogene extinction event.

<span class="mw-page-title-main">Tyrannosauridae</span> Family of dinosaurs

Tyrannosauridae is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous Tyrannosaurus. The exact number of genera is controversial, with some experts recognizing as few as three. All of these animals lived near the end of the Cretaceous Period and their fossils have been found only in North America and Asia.

<i>Tarbosaurus</i> Tyrannosaurid dinosaur genus from Late Cretaceous of Mongolia

Tarbosaurus is a genus of large tyrannosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 70 million years ago. It contains the single type species: Tarbosaurus bataar, which is known from the Nemegt Formation of Mongolia, with more fragmentary remains found further afield in the Subashi Formation of China. Tarbosaurus is represented by dozens of fossil specimens, including several complete skulls and skeletons. These remains have allowed studies focusing on its phylogeny, skull mechanics, and brain structure. Further fossil remains have been reported from other geologic formations of Asia, however, these remains are fragmentary and can not be confidentially assigned to Tarbosaurus or the type species.

<i>Daspletosaurus</i> Genus of tyrannosaurid dinosaur from Late Cretaceous period

Daspletosaurus is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 78 and 74.4 million years ago, during the Late Cretaceous Period. The genus Daspletosaurus contains three named species. Fossils of the earlier type species, D. torosus, have been found in Alberta, while fossils of a later species, D. horneri, have been found only in Montana. D. wilsoni has been suggested as an intermediate species between D. torosus and D. horneri that evolved through anagenesis, but this theory has been disputed by other researchers.

<i>Gorgosaurus</i> Genus of tyrannosaur dinosaur

Gorgosaurus is a genus of tyrannosaurid theropod dinosaur that lived in western North America during the Late Cretaceous Period (Campanian), between about 76.5 and 75 million years ago. Fossil remains have been found in the Canadian province of Alberta and the U.S. state of Montana. Paleontologists recognize only the type species, G. libratus, although other species have been erroneously referred to the genus.

<i>Aublysodon</i> Extinct genus of dinosaurs

Aublysodon is a genus of carnivorous dinosaurs known only from the Judith River Formation in Montana, which has been dated to the late Campanian age of the late Cretaceous period. The only currently recognized species, Aublysodon mirandus, was named by paleontologist Joseph Leidy in 1868. It is sometimes considered dubious now, because the type specimen consists only of an isolated premaxillary (front) tooth. Although this specimen is now lost, similar teeth have been found in many US states, western Canada, and Asia. These teeth almost certainly belong to juvenile tyrannosaurine tyrannosaurids, but most have not been identified to species level. However, it is likely that the type tooth belongs to one of the species in the genus Daspletosaurus, which was present in contemporary formations, and which matches specific details of the original tooth. The synapomorphies alleged to distinguish the Aublysodontinae, especially lack of serrations on premaxillary teeth could have been caused by tooth wear in life, postmortem abrasion, or digestion. Most other "aublysodontine"-type teeth may be from ontogenetic stages or sexual morphs of other tyrannosaurids.

<i>Alioramus</i> Tyrannosaurid theropod dinosaur genus from the Late Cretaceous period

Alioramus is a genus of tyrannosaurid theropod dinosaurs from the Late Cretaceous period of Asia. It currently contains two species. The type species, A. remotus is known from a partial skull and three foot bones recovered from the Mongolian Nemegt Formation, which was deposited in a humid floodplain about 70 million years ago. These remains were named and described by Soviet paleontologist Sergei Kurzanov in 1976. A second species, A. altai, known from a much more complete skeleton also from the Nemegt Formation, was named and described by Stephen L. Brusatte and colleagues in 2009. Its relationships to other tyrannosaurid genera were at first unclear, with some evidence supporting a hypothesis that Alioramus was closely related to the contemporary species Tarbosaurus bataar. However, the discovery of Qianzhousaurus indicates that it belongs to a distinct branch of tyrannosaurs, namely the tribe Alioramini.

<span class="mw-page-title-main">Tyrannosauroidea</span> Extinct superfamily of dinosaurs

Tyrannosauroidea is a superfamily of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent beginning in the Jurassic Period. By the end of the Cretaceous Period, tyrannosauroids were the dominant large predators in the Northern Hemisphere, culminating in the gigantic Tyrannosaurus. Fossils of tyrannosauroids have been recovered on what are now the continents of North America, Europe and Asia, with fragmentary remains possibly attributable to tyrannosaurs also known from South America and Australia.

<span class="mw-page-title-main">Albertosaurinae</span> Extinct subfamily of dinosaurs

Albertosaurines, or dinosaurs of the subfamily Albertosaurinae, lived in the Late Cretaceous of United States and Canada. The subfamily was first used by Philip J. Currie, Jørn H. Hurum, and Karol Sabath as a group of tyrannosaurid dinosaurs. It was originally defined as "(Albertosaurus + Gorgosaurus)", including only the two genera. The group is the sister clade to Tyrannosaurinae. In 2007, it was found that the group also contained Maleevosaurus, often synonymized with Tarbosaurus. However, this classification has not been accepted and Maleevosaurus is still considered a juvenile Tarbosaurus or Tyrannosaurus.

<i>Xiongguanlong</i> Extinct genus of dinosaurs

Xiongguanlong is an extinct genus of tyrannosauroid theropod from the Early Cretaceous period of what is now China. The type and only species is X. baimoensis. The generic name comes from Jiayuguan City and the Mandarin word "long" which means dragon. The specific epithet, "baimoensis" is a latinization of the Mandarin word for "white ghost" in reference to one of the geological features of the type locality.

<i>Bistahieversor</i> Extinct genus of dinosaurs

Bistahieversor, also known as the "Bisti Beast", is a genus of basal eutyrannosaurian theropod dinosaur. The genus contains only a single known species, B. sealeyi, described in 2010, from the Late Cretaceous of New Mexico. The holotype and a juvenile were found in the Hunter Wash Member of the Kirtland Formation, while other specimens came from the underlying Fossil Forest member of the Fruitland Formation. This dates Bistahieversor approximately 75.5 to 74.5 million years ago during the Campanian age, found in sediments spanning a million years.

<i>Teratophoneus</i> Extinct genus of dinosaurs

Teratophoneus is a genus of tyrannosaurine theropod dinosaur that lived during the late Campanian age of the Late Cretaceous period, in what is now Utah. It contains a single known species, T. curriei. It is known from an incomplete skull and postcranial skeleton recovered from the Kaiparowits Formation and was specifically named T. curriei in honor of famed paleontologist Philip J. Currie.

<i>Zhuchengtyrannus</i> Tyrannosaurid theropod dinosaur genus from Late Cretaceous period

Zhuchengtyrannus is a genus of tyrannosaurid theropod dinosaur known from the Campanian stage of the Late Cretaceous of Shandong Province, China. It belongs to the subfamily Tyrannosaurinae, and contains a single species, Zhuchengtyrannus magnus.

<i>Lythronax</i> Genus of tyrannosaurid dinosaur from the Late Cretaceous period

Lythronax is a genus of tyrannosaurid dinosaur that lived in North America around 81.9-81.5 million years ago during the Late Cretaceous period. The only known specimen was discovered in Utah in the Wahweap Formation of the Grand Staircase–Escalante National Monument in 2009, and it consists of a partial skull and skeleton. In 2013, it became the basis of the new genus and species Lythronax argestes; the generic name Lythronax means "gore king", and the specific name argestes originates from the Greek poet Homer's name for the wind from the southwest, in reference to the specimen's geographic provenance in North America.

<i>Qianzhousaurus</i> Extinct genus of dinosaurs

Qianzhousaurus is a genus of tyrannosaurid dinosaur that lived in Asia during the Maastrichtian age of the Late Cretaceous period. There is currently only one species named, the type species Qianzhousaurus sinensis, which is a member of the tribe Alioramini and most closely related to Alioramus, the only other known alioramin.

<span class="mw-page-title-main">Alioramini</span> Extinct clade of tyrannosaurid dinosaurs

Alioramini is a clade of long-snouted tyrannosaurine tyrannosaurids from the Late Cretaceous epoch. It includes the tyrannosaurid genera Alioramus and Qianzhousaurus. Although tyrannosaurids are known from a variety of places around the globe, alioramins are currently restricted to Asia in mostly Maastrichtian strata. Many of the fossils attributed to Alioramini are not from fully developed individuals.

<span class="mw-page-title-main">Timeline of tyrannosaur research</span>

This timeline of tyrannosaur research is a chronological listing of events in the history of paleontology focused on the tyrannosaurs, a group of predatory theropod dinosaurs that began as small, long-armed bird-like creatures with elaborate cranial ornamentation but achieved apex predator status during the Late Cretaceous as their arms shrank and body size expanded. Although formally trained scientists did not begin to study tyrannosaur fossils until the mid-19th century, these remains may have been discovered by Native Americans and interpreted through a mythological lens. The Montana Crow tradition about thunder birds with two claws on their feet may have been inspired by isolated tyrannosaurid forelimbs found locally. Other legends possibly inspired by tyrannosaur remains include Cheyenne stories about a mythical creature called the Ahke, and Delaware stories about smoking the bones of ancient monsters to have wishes granted.

<i>Dynamoterror</i> Extinct genus of dinosaurs

Dynamoterror is a monospecific genus of tyrannosaurid dinosaur from New Mexico that lived during the Late Cretaceous in what is now the upper Allison Member of the Menefee Formation. The type and only species, Dynamoterror dynastes, is known from a subadult or adult individual about 9 metres long with an incomplete associated skeleton. It was named in 2018 by Andrew T. McDonald, Douglas G. Wolfe and Alton C. Dooley, Jr. Dynamoterror was closely related to Teratophoneus and Lythronax.

<i>Thanatotheristes</i> Daspletosaurin tyrannosaurid of the mid-Campanian

Thanatotheristes is a genus of tyrannosaurid dinosaur from the Late Cretaceous of Laramidia, approximately 80.1-79.5 Ma. Thanatotheristes contains only one species, T. degrootorum. Fossils of this taxon are found in the Foremost Formation of Alberta, Canada, coexisting with medium-sized ceratopsids like Xenoceratops foremostensis and small pachycephalosaurids like Colepiocephale lambei.

<span class="mw-page-title-main">Daspletosaurini</span> Extinct clade of tyrannosaurid dinosaurs

Daspletosaurini is an extinct clade of tyrannosaurine dinosaurs that lived in Laramidia during the Late Cretaceous period. It consists of two genera: Daspletosaurus and Thanatotheristes. Four species have been described in the two genera, namely Daspletosaurus torosus, Daspletosaurus horneri, Daspletosaurus wilsoni, and Thanatotheristes degrootorum. At one point all of them were assigned as specimens of D. torosus, but several papers since 2017 have found them to represent distinct species. Some researchers found anagenesis in the group, whether contained in a daspletosaurin clade or paraphyletic in respect to the lineage of tyrannosaurines leading up to Tyrannosaurus, but a 2023 study refuted this theory on the basis of morphological and stratigraphical data.

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. ↑ Zheng, Wenjie; Jin, Xingsheng; Xie, Junfang; Du, Tianming (2024-07-25). "The first deep-snouted tyrannosaur from Upper Cretaceous Ganzhou City of southeastern China". Scientific Reports . 14 (1): 16276. Bibcode:2024NatSR..1416276Z. doi:10.1038/s41598-024-66278-5. ISSN   2045-2322. PMC   11272791 . PMID   39054316.
  3. 1 2 3 4 Scherer, Charlie Roger; Voiculescu-Holvad, Christian (2023-11-28). "Re-analysis of a dataset refutes claims of anagenesis within Tyrannosaurus-line tyrannosaurines (Theropoda, Tyrannosauridae)". Cretaceous Research . 155: 105780. doi: 10.1016/j.cretres.2023.105780 . ISSN   0195-6671.
  4. ↑ Rivera-Sylva, Héctor E.; Longrich, Nicholas R. (2024). "A New Tyrant Dinosaur from the Late Campanian of Mexico Reveals a Tribe of Southern Tyrannosaurs". Fossil Studies. 2 (4): 245–272. doi: 10.3390/fossils2040012 .
  5. ↑ 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.
  6. ↑ 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.
  7. ↑ Maleev, Evgeny A. (1955). "Giant carnivorous dinosaurs of Mongolia". Doklady Akademii Nauk SSSR . 104 (4): 634–637.
  8. ↑ 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.
  9. ↑ Rozhdestvensky, Anatoly K. (1965). "Growth changes in Asian dinosaurs and some problems of their taxonomy". Paleontological Journal. 3: 95–109.
  10. ↑ Russell, Dale A. (1970). "Tyrannosaurs from the Late Cretaceous of western Canada". National Museum of Natural Sciences Publications in Paleontology. 1: 1–34.
  11. ↑ 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.
  12. ↑ Dong Zhiming (1977). "On the dinosaurian remains from Turpan, Xinjiang". Vertebrata PalAsiatica (in Chinese). 15: 59–66.
  13. ↑ 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.
  14. ↑ 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.
  15. ↑ 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.
  16. ↑ Olshevsky, G. (1995). "The origin and evolution of the tyrannosaurids". Kyoryugaku Saizensen [Dino Frontline]. 9–10: 92–119.
  17. ↑ Carr, Thomas D. (2022-11-25). "A reappraisal of tyrannosauroid fossils from the Iren Dabasu Formation (Coniacian–Campanian), Inner Mongolia, People's Republic of China". Journal of Vertebrate Paleontology. 42 (5). Bibcode:2022JVPal..42E9817C. doi: 10.1080/02724634.2023.2199817 . ISSN   0272-4634.
  18. ↑ Carr, Thomas D. (1999). "Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)". Journal of Vertebrate Paleontology. 19 (3): 497–520. Bibcode:1999JVPal..19..497C. doi:10.1080/02724634.1999.10011161. S2CID   83744433.
  19. ↑ Currie, Philip J.; Dong Zhiming (2001). "New information on Shanshanosaurus huoyanshanensis, a juvenile tyrannosaurid (Theropoda, Dinosauria) from the Late Cretaceous of China" (PDF). Canadian Journal of Earth Sciences. 38 (12): 1729–1737. Bibcode:2001CaJES..38.1729C. doi:10.1139/cjes-38-12-1729.
  20. 1 2 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...621376F. doi: 10.1371/journal.pone.0021376 . PMC   3126816 . PMID   21738646.
  21. ↑ 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.
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