Tropeognathus

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Tropeognathus
Temporal range: Early Cretaceous (Albian), 112  Ma
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Tropeognathus mesembrinus MN 01.jpg
Reconstructed skeleton, National Museum of Brazil
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order:† Pterosauria
Suborder:† Pterodactyloidea
Family:† Anhangueridae
Subfamily:† Tropeognathinae
Genus:† Tropeognathus
Wellnhofer, 1987
Type species
†Tropeognathus mesembrinus
Wellnhofer, 1987
Synonyms

Tropeognathus (meaning "keel jaw") is a genus of large pterosaurs from the late Early Cretaceous of South America. This genus is considered to be a member of the family Anhangueridae, however, several studies have also recovered it within another family called Ornithocheiridae. Both of these families are diverse groups of pterosaurs known for their keel-tipped snouts and large size. Tropeognathus is regarded as the largest pterosaur found in the Southern Hemisphere, only rivaled by the huge azhdarchids. [1] The type and only species is Tropeognathus mesembrinus. Fossil remains of Tropeognathus have been recovered from the Romualdo Formation, which is a Lagerstätte located in the Santana Group of the Araripe Basin in northeastern Brazil. [2]

Discovery and naming

Holotype specimen of Tropeognathus mesembrinus (BSP 1987 I 46) in lateral (A1), palatal (A2), and anterior (A3) views Tropeognathus.png
Holotype specimen of Tropeognathus mesembrinus (BSP 1987 I 46) in lateral (A1), palatal (A2), and anterior (A3) views

In the 1980s the German paleontology museum Bayerische Staatssammlung für Paläontologie und historische Geologie in Munich acquired a pterosaur skull from Brazilian fossil dealers that had probably been found in Ceará, in the geological group called the Santana Group, which is located in the Araripe Basin (Chapada do Araripe) of Brazil. [3] In 1987, it was named and described as the type species Tropeognathus mesembrinus by Peter Wellnhofer. The generic name is derived from Greek τρόπις, tropis, "keel", and γνάθος, gnathos, "jaw". The specific name is derived from Koine mesembrinos, "of the noontide", simplied as "southern", in reference to the provenance from the Southern hemisphere. [3]

The holotype, BSP 1987 I 46, was discovered in a layer of the Romualdo Formation within the Santana Group, dating to the latest Aptian and earliest Albian stages. Along with the holotype, several other pterosaur specimens were found in the fossil site, these specimens however, were referred to genera such as Anhanguera and Cearadactylus . [4] [5] The uncovered holotype consists of a skull with lower jaws. A second specimen was referred by André Jacques Veldmeijer in 2002: SMNS 56994, which consists of a partial mandible. [6] In 2013, Brazilian paleontologist Alexander Kellner referred a third, larger, specimen: MN 6594-1, a skeleton with skull, with extensive elements of all body parts, except the tail and the lower hindlimbs. [7]

After Tropeognathus mesembrinus was named by Peter Wellnhofer in 1987, other researchers tended to consider it part of several other genera, leading to an enormous taxonomic confusion. [8] It was considered an Anhanguera mesembrinus by Alexander Kellner in 1989, [9] a Criorhynchus mesembrinus by Veldmeijer in 1998 and a Coloborhynchus mesembrinus by Michael Fastnacht in 2001. [10] [11] [7] Later the same year, David Unwin referred the Tropeognathus material to Ornithocheirus simus, making Tropeognathus mesembrinus a junior synonym, though he had reinstated a Ornithocheirus mesembrinus in 2003. [12] [13] In 2006, Veldmeijer accepted that Tropeognathus and Ornithocheirus were cogeneric, but rejected O. simus as the type species of Ornithocheirus in favor of O. compressirostris, which was named as Lonchodectes by Unwin due to an analysis by English paleontologist Reginald Walter Hooley in 1914. [14] [15] This made Veldmeijer use the names Criorhynchus simus and Criorhynchus mesembrinus instead. [2] In 2013 however, Taissa Rodrigues and Alexander Kellner concluded Tropeognathus to be valid, and containing only T. mesembrinus, the type species. [7]

Back in 1987, Wellnhofer had named a second species called Tropeognathus robustus, based on specimen BSP 1987 I 47, which is a more robust lower jaw. [3] In 2013 however, T. robustus was considered as a species of Anhanguera, resulting in an Anhanguera robustus. [7]

Description

Size

Size comparison of Tropeognathus mesembrinus Tropeognathus mesembrinus size chart.png
Size comparison of Tropeognathus mesembrinus
Restoration of three flying individuals TropeognathusDB22.jpg
Restoration of three flying individuals

Tropeognathus is known to have reached wingspans of about 8.26 meters (27.1 ft), as can be inferred from the impressive size of the specimen MN 6594-1. [1] The maximum wingspan estimate for Tropeognathus reaches 8.70 meters (28.5 ft), [1] making it slightly larger than the average estimate for the genus, though much larger than other close relatives such as Ornithocheirus and Coloborhynchus, which are typically estimated to be between 4.5 to 6.1 meters (15 to 20 ft). [7] A skull unearthed belonging to the related Coloborhynchus likely measured 75 centimeters (30 in), which led to a wingspan estimate of 7 meters (23 ft) respectively: larger than the average estimates for this genus, but still shorter than that of Tropeognathus. This specimen however, was concluded to belong to another genera, though in several studies, some paleontologists consider it under the species Coloborhynchus capito, which was originally called Ornithocheirus capito by the British paleontologist Harry Govier Seeley back in 1870. [16] [17]

Skull and crests

Restored fossil skull of Tropeognathus; note the prominent, keel-like snout-crests. Tropeognathus.JPG
Restored fossil skull of Tropeognathus; note the prominent, keel-like snout-crests.
Skull comparison of different anhanguerians, notice Tropeognathus (E and F) with a well-developed "keeled" crest Anhangueria skulls.jpg
Skull comparison of different anhanguerians, notice Tropeognathus (E and F) with a well-developed "keeled" crest

The skull of Tropeognathus bore a distinctively convex, "keeled" crests on the lower end of its snout, with an opposing, smaller mass on the underside of the lower jaws. This structure was prominent, well-developed, and relatively large in Tropeognathus (especially in males); however, by comparison, these crests were somewhat weakly-formed as opposed to the thicker skull crests of other pterosaurs, such as Ornithocheirus. [2] The upper crests arose from the snout tip and extended back to the fenestra nasoantorbitalis, the large opening in the skull side. An additional, smaller crest projected down from the lower jaws at their symphysis ("chin" area). [14] The similar anhanguerid Anhanguera possessed jaws that were tapered in width, but expanded into a broad, spoon-shaped rosette at the tip, which differed from Tropeognathus for having a narrower appearance. [18] The jaws can be distinguished from its relatives by a few differences in the crest: unlike its close relatives Coloborhynchus and Ornithocheirus, the crest on the upper jaw of Tropeognathus was more prominent and much larger, and therefore resulting in a broader skull. [1]

Vertebrae

The first five dorsal vertebrae of Tropeognathus are fused into a notarium, with five sacral vertebrae fused into a synsacrum, and the third and fourth sacral vertebrae are keeled within. The front blade of the ilium is strongly directed upwards, resulting in a narrow structure. [1]

Classification

Palate of Tropeognathus (B) compared to the palates of Ferrodraco (A) and Siroccopteryx (C), all of them are in occlusal view Tropeognathine palates.png
Palate of Tropeognathus (B) compared to the palates of Ferrodraco (A) and Siroccopteryx (C), all of them are in occlusal view

In 1987, Wellnhofer assigned Tropeognathus to a Tropeognathidae. [3] This concept was not adopted by other workers; several researchers place Tropeognathus mesembrinus in the Anhangueridae, along with Anhanguera, while other cladistic analyses place Tropeognathus within the Ornithocheiridae as a basal member, meaning that it was more closely related to Ornithocheirus than Anhanguera. This concept is mostly used by the European colleagues, who prefer to use the Ornithocheiridae as the most inclusive group rather than the Anhangueridae. [13] A topology made by Andres and Myers in 2013 placed Tropeognathus within the family Ornithocheiridae in a more basal position than Ornithocheirus, and the family itself is placed within the more inclusive clade Ornithocheirae. [19] However, many subsequent analyses made in 2019 and 2020 have recovered Tropeognathus within the family Anhangueridae, [20] [21] [22] with a specific one by Borja Holgado and Rodrigo Pêgas in 2020, placing Tropeognathus more specifically within the subfamily Tropeognathinae, sister taxon to Siroccopteryx . [23]

Hand puppet from Walking With Dinosaurs, Oxford Museum Pterosaur exhibit.jpg
Hand puppet from Walking With Dinosaurs, Oxford Museum

Tropeognathus mesembrinus was the subject of an entire episode of the award-winning BBC television program Walking with Dinosaurs (which used the first name of its cousin Ornithocheirus but was incorrectly named as a species of it, as Ornithocheirus mesembrinus). [14] In Walking with Dinosaurs: A Natural History, a companion book to the series, it was claimed that several large bone fragments from the Santana Group (known as Santana Formation in the book) of Brazil had indicated that O. mesembrinus may have had a wingspan reaching 12 meters (39 ft) and a weight of 100 kilograms (220 lb), making it one of the largest known pterosaurs. [25] However, the largest definite Ornithocheirus mesembrinus specimens described at the time measured 6 meters (20 ft), in terms of wingspan. [8] The specimens which the producers of the program used to justify such a large size estimate were described in 2012, and were under study by Dave Martill and David Unwin at the time of Walking With Dinosaurs' production. The final description of the remains found a maximum estimated wingspan of 8.70 meters (28.5 ft) for this large specimen. [1] Unwin stated that he did not believe the higher estimate used by the BBC was likely, and that the producers likely chose the highest possible estimate because it was more "spectacular." [26] Nevertheless, specimen MN 6594-V in 2013 was, at its degree of completeness, the largest known pterosaur individual. [1]

See also

Related Research Articles

<i>Cearadactylus</i> Genus of anhanguerid pterosaur from the Early Cretaceous

Cearadactylus is a genus of large anhanguerid pterodactyloid pterosaur from the Romualdo Formation of Brazil, South America. Fossil remains of Cearadactylus dated back to the Albian stage of the Early Cretaceous period, about 112 million years ago. The only known species is C. atrox, described and named in 1985 by Giuseppe Leonardi and Guido Borgomanero. The name refers to the Brazilian state Ceará, and combines this with Greek daktylos, "finger", a reference to the wing finger of pterosaurs. The Latin atrox means "frightful", a reference to the fearsome dentition of the species.

<i>Ornithocheirus</i> Genus of ornithocheirid pterosaur from the Early Cretaceous

Ornithocheirus is a pterosaur genus known from fragmentary fossil remains uncovered from sediments in the United Kingdom and possibly Morocco.

<i>Thalassodromeus</i> Genus of azhdarchoid pterosaur from the Early Cretaceous

Thalassodromeus is a genus of pterosaur that lived in what is now Brazil during the Early Cretaceous period, about a hundred million years ago. The original skull, discovered in 1983 in the Araripe Basin of northeastern Brazil, was collected in several pieces. In 2002, the skull was made the holotype specimen of Thalassodromeus sethi by palaeontologists Alexander Kellner and Diogenes de Almeida Campos. The generic name means "sea runner", and the specific name refers to the Egyptian god Seth due to its crest being supposedly reminiscent of Seth's crown. Other scholars have pointed out that the crest was instead similar to the crown of Amon. A jaw tip was assigned to T. sethi in 2005, became the basis of the new genus Banguela in 2014, and assigned back to Thalassodromeus as the species T. oberlii in 2018. Another species was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell.

Anhanguera is a genus of pterodactyloid pterosaur known from the Early Cretaceous Romualdo Formation of Brazil and the Late Cretaceous Kem Kem Group of Morocco. This pterosaur is closely related to Ornithocheirus, but belongs in the family Anhangueridae. The generic name comes from the Tupi words añanga, meaning "spirit protector of the animals" + wera "bygone".

Araripesaurus is a genus a pterosaur belonging to the suborder Pterodactyloidea, it was discovered in the Romualdo Formation of the Santana Group in northeastern Brazil, which dates back to the Aptian and Albian of the Early Cretaceous. The type species is A. castilhoi.

Brasileodactylus a genus of pterosaur from the Aptian-age lower Santana formation of Chapada do Araripe, Ceará, Brazil.

<i>Siroccopteryx</i> Genus of anhanguerid pterosaur from the Early Cretaceous

Siroccopteryx is an extinct genus of anhanguerid pterodactyloid pterosaur, known from middle Cretaceous sediments in modern-day Morocco. Some researchers, such as David M. Unwin, consider the genus a junior synonym of Coloborhynchus.

<i>Coloborhynchus</i> Genus of anhanguerid pterosaur from the Cretaceous period

Coloborhynchus is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. Coloborhynchus is known from the Lower Cretaceous of England, and depending on which species are included, possibly the Albian and Cenomanian ages as well. Coloborhynchus was once thought to be the largest known toothed pterosaur, however, a specimen of the closely related Tropeognathus is now thought to have had a larger wingspan.

<i>Santanadactylus</i> Genus of pteranodontoid pterosaur from the Early Cretaceous

Santanadactylus was a genus of pterodactyloid pterosaur from the Albian-age Romualdo Member of the Upper Cretaceous Santana Formation, of Barra do Jardim, Araripe Plateau, Ceará State, Brazil. Four species have been named, but today are no considered congeneric with each other. It was a rather large pterosaur.

<span class="mw-page-title-main">Ornithocheiridae</span> Family of ornithocheiran pterosaurs

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<i>Uktenadactylus</i> Genus of anhanguerid pterosaur from the Early Cretaceous

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<span class="mw-page-title-main">Anhangueridae</span> Family of anhanguerian pterosaurs

Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a lateral expansion in the distal rostrum. The same study presented a cladistic analysis, for which an "agreement subtree" was calculated. The Anhangueridae was found to be sister taxon to the large crested Tropeognathus.

<span class="mw-page-title-main">Pteranodontoidea</span> Clade of ornithocheiroid pterosaurs from the Cretaceous period

Pteranodontoidea is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous of Asia, Africa, Europe, North America and South America. It was named by Alexander Wilhelm Armin Kellner in 1996. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Anhanguera, Pteranodon and all its descendants. The clade Ornithocheiroidea is sometimes considered to be the senior synonym of Pteranodontoidea, however it depends on its definition. Brian Andres in his analyses, converts Ornithocheiroidea using the definition of Kellner (2003) to avoid this synonymy.

<span class="mw-page-title-main">Ornithocheirae</span> Clade of pteranodontoid pterosaurs

Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous of Asia, Europe, North America and South America. It was named by Harry Seeley in 1870 as a family that contains Ornithocheirus and its relatives. The name was emended to Ornithocheiridae, to match the requirements of the ICZN Code that a family-ranked clade should end with an -idae suffix. Brian Andres (2010) in his review of pterosaur phylogeny, defined the name Ornithocheirae phylogenetically, as a node-based taxon consisting of the last common ancestor of Anhanguera and Ornithocheirus and all its descendants. Thus Ornithocheirae is defined to include two families, the Anhangueridae and the Ornithocheiridae, following the opinion of Alexander Kellner and Andres that these families should not be synonymized based on their original phylogenetical definitions. However, subsequent studies in 2019 have found Ornithocheirae to be a more inclusive group containing both Anhangueria and Targaryendraconia.

<i>Cimoliopterus</i> Genus of pterosaur from the Cretaceous period

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<i>Camposipterus</i> Genus of ornithocheiran pterosaur from the Early Cretaceous

Camposipterus is a genus of pterodactyloid pterosaur from the Early Cretaceous of England. Fossil remains of Camposipterus dated back to the Early Cretaceous, about 112 million years ago.

<span class="mw-page-title-main">Anhangueria</span> Clade of ornithocheiran pterosaurs from the Cretaceous period

Anhangueria is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 92.5 million years ago. Anhangueria was named by paleontologists Taissa Rodrigues and Alexander Kellner in a review of Ornithocheirus species in 2013, they defined the clade as a branch-based taxon consisting of all pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri.

<i>Maaradactylus</i> Genus of anhanguerid pterosaur from the Early Cretaceous

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<span class="mw-page-title-main">Ornithocheiromorpha</span> Clade of pteranodontoid pterosaurs

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<i>Nicorhynchus</i> Genus of anhanguerid pterosaur from the Cretaceous period

Nicorhynchus is a genus of anhanguerid pterosaur from the Cretaceous period. It contains two species, the type species, N. capito, from the Cambridge Greensand of England, and N. fluviferox from the Kem Kem Group of Morocco. These species were previously assigned to Coloborhynchus.

References

  1. 1 2 3 4 5 6 7 Kellner, A. W. A.; Campos, D. A.; Sayão, J. M.; Saraiva, A. N. A. F.; Rodrigues, T.; Oliveira, G.; Cruz, L. A.; Costa, F. R.; Silva, H. P.; Ferreira, J. S. (2013). "The largest flying reptile from Gondwana: A new specimen of Tropeognathus cf. T. Mesembrinus Wellnhofer, 1987 (Pterodactyloidea, Anhangueridae) and other large pterosaurs from the Romualdo Formation, Lower Cretaceous, Brazil". Anais da Academia Brasileira de Ciências. 85 (1): 113–135. doi: 10.1590/S0001-37652013000100009 . PMID   23538956.
  2. 1 2 3 Veldmeijer, A.J. (2006). "Toothed pterosaurs from the Santana Formation (Cretaceous; Aptian-Albian) of northeastern Brazil. A reappraisal on the basis of newly described material Archived 2012-03-17 at the Wayback Machine ." Tekst. – Proefschrift Universiteit Utrecht.
  3. 1 2 3 4 Peter Wellnhofer, 1987, "New crested pterosaurs from the Lower Cretaceous of Brazil", Mitteilungen der Bayerische Staatssammlung für Paläontologie und historische Geologie27: 175–186; Muenchen
  4. ↑ Pinheiro, F.L.; Rodrigues, Taissa (2017). "Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?". PeerJ. 5: e3285. doi: 10.7717/peerj.3285 . PMC   5420195 . PMID   28484676.
  5. ↑ Leonardi, G. & Borgomanero, G. (1985). "Cearadactylus atrox nov. gen., nov. sp.: novo Pterosauria (Pterodactyloidea) da Chapada do Araripe, Ceara, Brasil." Resumos dos communicaçoes VIII Congresso bras. de Paleontologia e Stratigrafia, 27: 75–80.
  6. ↑ Veldmeijer, A.J. (2002). "Pterosaurs from the Lower Cretaceous of Brazil in the Stuttgart collection". Stuttgarter Beiträge zur Naturkunde, Serie B (Geologie und Paläontologie). 327: 1–27.
  7. 1 2 3 4 5 Rodrigues, T.; Kellner, A. (2013). "Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England". ZooKeys (308): 1–112. Bibcode:2013ZooK..308....1R. doi: 10.3897/zookeys.308.5559 . PMC   3689139 . PMID   23794925.
  8. 1 2 Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs. New York: Barnes and Noble Books. pp. 124. ISBN   0-7607-0154-7.
  9. ↑ Kellner, A.W.A. (1989). "A new Edentate Pterosaur of the lower Cretaceous from the Araripe Basin, Northeast Brazil". Anais da Academia Brasileira de Ciências. 61: 439–446. S2CID   89420181.
  10. ↑ Veldmeijer, A.J. (1998). "Pterosaurs from the Lower Cretaceous of Brazil in the Stuttgart Collection". Geoscience and Engineering. 327: 1–27.
  11. ↑ Fastnacht, M (2001). "First record of Coloborhynchus (Pterosauria) from the Santana Formation (Lower Cretaceous) of the Chapada do Araripe of Brazil". Paläontologische Zeitschrift. 75 (1): 23–36. Bibcode:2001PalZ...75...23F. doi:10.1007/bf03022595. S2CID   128410270.
  12. ↑ Unwin, D.M., 2001, "An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England", Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe4: 189–221
  13. 1 2 Unwin, D. M. (2003). "On the phylogeny and evolutionary history of pterosaurs". Geological Society, London, Special Publications. 217 (1): 139–190. Bibcode:2003GSLSP.217..139U. CiteSeerX   10.1.1.924.5957 . doi:10.1144/GSL.SP.2003.217.01.11. S2CID   86710955.
  14. 1 2 3 Unwin, David M. (2006). The Pterosaurs: From Deep Time. New York: Pi Press. p. 246. ISBN   0-13-146308-X.
  15. ↑ Hooley, Reginald Walter (1914). "On the Ornithosaurian genus Ornithocheirus, with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge". Annals and Magazine of Natural History. 13 (78): 529–557. doi:10.1080/00222931408693521. ISSN   0374-5481.
  16. ↑ Martill, D.M. and Unwin, D.M. (2011). "The world's largest toothed pterosaur, NHMUK R481, an incomplete rostrum of Coloborhynchus capito (Seeley 1870) from the Cambridge Greensand of England." Cretaceous Research, (advance online publication). doi : 10.1016/j.cretres.2011.09.003
  17. ↑ Seeley, H.G. (1870). The Ornithosauria: an Elementary Study of the Bones of Pterodactyles. Cambridge, 130 pp.
  18. ↑ Kellner, A.W.A. and Tomida, Y. (2000). "Description of a new species of Anhanguera (Pterodactyloidea) with comments on the pterosaur fauna from the Santana Formation (Aptian–Albian), northeastern Brazil." Tokyo, National Science Museum (National Science Museum Monographs, 17).
  19. 1 2 Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 1. doi:10.1017/S1755691013000303. S2CID   84617119.
  20. ↑ Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", Scientific Reports9: 4940 doi : 10.1038/s41598-019-41280-4
  21. ↑ Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019). "First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity". Scientific Reports. 9 (1): 17875. Bibcode:2019NatSR...917875K. doi: 10.1038/s41598-019-54042-z . PMC   6884559 . PMID   31784545.
  22. ↑ Pêgas, R.V., Holgado, B., Leal, M.E.C., 2019. "Targaryendraco wiedenrothi gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids", Historical Biology, 1–15. doi:10.1080/08912963.2019.1690482
  23. 1 2 Holgado, B.; Pêgas, R.V. (2020). "A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae". Acta Palaeontologica Polonica. 65. doi: 10.4202/app.00751.2020 .
  24. ↑ "Dinosaurs in the Museum" (PDF). Retrieved February 15, 2024.
  25. ↑ Haines, T., 1999, "Walking with Dinosaurs": A Natural History, BBC Books, p. 158
  26. ↑ Bredow, H.P. (April 18, 2000). "Re: WWD non-dino questions". Archived from the original on April 4, 2016. Retrieved January 20, 2011.
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