Thalassodromidae

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Thalassodromids
Temporal range: Early-Late Cretaceous, 112–88  Ma
DSC 0042.JPG
Replica skeleton of Tupuxuara leonardii
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order:† Pterosauria
Suborder:† Pterodactyloidea
Clade:† Azhdarchoidea
Family:† Thalassodromidae
Witton, 2009
Type species
† Thalassodromeus sethi
Kellner & Campos, 2002
Genera
Synonyms
  • Thalassodrominae
    Kellner, 2007
  • Tupuxuaridae
    Martill, Bechly & Heads, 2007

Thalassodromidae (meaning "sea runners", due to previous misconceptions of skimming behavior; they are now thought to be terrestrial predators) is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional members come from Brazil, which include the type genus Thalassodromeus , as well as Tupuxuara . Additionally, the recently named genus Kariridraco , also from Brazil, is another undisputed thalassodromid. Proposals of additional thalassodromid genera have proved more controversial.

The classification of Thalassodromidae is quite controversial and disputed. It was initially denominated Thalassodrominae, as a subfamily within the group Tapejaridae. However, opposing studies regarding its placement have argued that its members were more closely related to azhdarchids and dsungaripterids, while also elevating it to family level. Though the relationship with tapejarids remains supported by many studies, it has been proposed that it is preferable to retain them as a distinct family for consistency of communication.

Classification

Life reconstruction of thalassodromid Kariridraco Kariridraco by Julia d'Oliveira.jpg
Life reconstruction of thalassodromid Kariridraco

The classification of thalassodromids is controversial. Its initial members included Thalassodromeus and Tupuxuara , which were assigned initially to the family Tapejaridae by Brazilian paleontologists Alexander Kellner and Diógenes de Almeida Campos. In 2007, the subfamily Thalassodrominae was coined by Kellner to group both members within Tapejaridae. [1]

However, a conflicting phylogenetic model emerged arguing that both Thalassodromeus and Tupuxuara were more closely related to the family Azhdarchidae instead of the tapejarids. As early as 2003, paleontologist David Unwin created the group Neoazhdarchia to contain Tupuxuara and Azhdarchidae. [2] This arrangement would later be supported by British paleontologists David Martill and Darren Naish, who deemed Tapejaridae to be paraphyletic (unnatural), and found both Thalassodromeus and Tupuxuara as sister taxa to Azhdarchidae. [3] In 2008, Chinese paleontologist Lü Junchang and colleagues would use the term "Tupuxuaridae" to include both genera in their phylogenetic analysis, finding the group nested within Neoazhdarchia. [4] In 2009, British paleontologist Mark Witton also concurred with the placement of Thalassodromeus and Tupuxuara within Neoazhdarchia. However, he noted that the term Thalassodrominae was created before Tupuxuaridae, so therefore it had naming priority. He elevated Thalassodrominae to family level to satisfy the hierarchy within Neoazhdarchia, thus creating the denomination Thalassodromidae. [5] Phylogenetic analyses by Brian Andres in 2014 and 2021 would support this model, finding a relationship between Thalassodromidae and Dsungaripteridae within Neoazhdarchia, a conclusion also found by a 2018 study. [6] [7] [8] [9]

Many studies, however, have retained the original model of Thalassodrominae as a subfamily of Tapejaridae. In 2011 a study Brazilian paleontologist Felipe Pinheiro and colleagues retained the conception and expanded it to include Chaoyangopterinae, considered by other studies to be a distinct family. [10] Studides by Rodrigo Pêgas and colleagues, [11] [12] Gabriela Cerquiera and colleagues, [13] [14] as well as Kellner himself have continued to followed Kellner's original model, into the 2020s, [15] [16] amongst others. [17] [18] Despite continuing to favour the close relationship of Thalassodromeus and Tupuxuara with Tapejara and its relatives, Pêgas and colleagues began to advocate for the use of Thalassodromidae rather than Thalassodrominae for sake of consistency between studies that do and do not find this relationship. Under this nomenclatural model, they are both families within Tapejaromorpha. [19] Subsequent studies by Pêgas and colleagues would follow this change, [12] [20] though others continue to use the traditional scope of Tapejaridae. [14] [16]

Reconstructed skull of Thalassodromeus; disagreement exists over whether the jaw would downturn as shown Thalassodromeus skull.png
Reconstructed skull of Thalassodromeus ; disagreement exists over whether the jaw would downturn as shown

In 2018, paleontologist Nicholas Longrich and colleagues recovered the pterosaurs Aerotitan and Alanqa as thalassodromids, but noted that their fossil remains of are fragmentary, so this assignment was only tentative. [9] A 2021 study by Brian Andres also found Alanqa as a thalassodromid, as well as Argentinadraco , Leptostomia , and Xericeps within Thalassodromidae. He redefined the term Thalassodrominae for all taxa closer to Thalassodromeus than to Tupuxuara, which included all four of this genera. [8] Other studies would not corroborate these results. In a 2022 study, Pêgas and colleagues re-evaluated Aerotitan and rejected its identity as a thalassodromid. [21] Other studies have supported this conclusion. [16] [12] The 2022 study also reinterpreted the holotype of Alanqa as a lower jaw as opposed to an upper one, and found it did not resemble those of thalassodromids. They concluded it to be more closely related to azhdarchids and named a new family Alanqidae for it. [21] A 2023 study by Roy Smith and all authors of the 2018 study would agree with this interpretation, rejecting Alanqa as a thalassodromid (though considering it an azhdarchid rather than an alanqid). They also expressed skepticism at the result of Leptosomia and Xericeps as thalassodromids, noting their fragmentary nature and concluding it was difficult to be certain of their position within Azhdarchoidea. [22] Subsequent studies have placed these two genera and Argentinadraco in Alanqidae, Azhdarchidae, or Chaoyangopteridae. [12] [16] [23]

Related Research Articles

<span class="mw-page-title-main">Azhdarchidae</span> Family of large azhdarchoid pterosaurs

Azhdarchidae is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cretaceous as well. Azhdarchids are mainly known for including some of the largest flying animals discovered, but smaller cat-size members have also been found. Originally considered a sub-family of Pteranodontidae, Nesov (1984) named the Azhdarchinae to include the pterosaurs Azhdarcho, Quetzalcoatlus, and Titanopteryx. They were among the last known surviving members of the pterosaurs, and were a rather successful group with a worldwide distribution. Previously it was thought that by the end of the Cretaceous, most pterosaur families except for the Azhdarchidae disappeared from the fossil record, but recent studies indicate a wealth of pterosaurian fauna, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms.

<i>Tupuxuara</i> Genus of thalassodromid pterosaur from the Early Cretaceous

Tupuxuara is a genus of thalassodromid pterosaur that lived during the Albian age of the Early Cretaceous, about 112 million years ago. Its remains were found in what is now the Romualdo Formation of the Santana Group in Brazil. Tupuxuara was named in 1988 by paleontologists Alexander Kellner and Diógenes de Almeida Campos. The name Tupuxuara means "familiar spirit" referring to a familiar spirit in the mythology of the Tupi people in Brazil. Two species have been named, T. longicristatus, the type species, and T. leonardii. An additional species has been named in 2013, T. deliradamus. However, the validity of this species has been put into question and it may not even belong to Tupuxuara.

<i>Thalassodromeus</i> Genus of thalassodromid 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 2015, and assigned back to Thalassodromeus as the species T. oberlii in 2018, though other researchers consider it a valid genus. Another species was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell.

<i>Arambourgiania</i> Genus of large azhdarchid pterosaur from the Late Cretaceous

Arambourgiania is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous, in what is now Jordan. Additional fossil remains from the United States and Morocco have also been found, but their assignment to Arambourgiania is only tentative. The original specimen was discovered in the 1940s by a railway worker near Russeifa, Jordan. After examination by paleontologist Camille Arambourg, a new species was named in 1959, Titanopteryx philadelphiae. The generic name means "titan wing", as the fossil was initially misidentified as a huge wing metacarpal, while the specific name refers to the ancient name of Amman, Philadelphia. The genus "Titanopteryx" would later be problematic, as it had already been taken by a fly. Because of this, paleontologist Lev Nessov in 1989 named a new genus, Arambourgiania, in honor of Arambourg. The new species was now known as Arambourgiania philadelphiae.

<span class="mw-page-title-main">Tapejaridae</span> Family of azhdarchoid pterosaurs from the Cretaceous period

Tapejaridae is a family of azhdarchoid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primitive genera were found in China, indicating that the family has an Asian origin.

<i>Bennettazhia</i> Genus of tapejaromorph pterosaur from the Early Cretaceous

Bennettazhia is a genus of tapejaromorph pterosaur which lived during the Albian stage of the Early Cretaceous from what is now the Hudspeth Formation of the state of Oregon in the United States. Although originally identified as a species of the pteranodontoid pterosaur Pteranodon, Bennettazhia is now thought to have been a different animal. The type and only species is B. oregonensis.

<i>Eopteranodon</i> Genus of tapejarid pterosaur

Eopteranodon is a genus of tapejarid pterosaur from the Aptian-age Lower Cretaceous Yixian Formation of Beipiao City, Liaoning, China. The genus was named in 2005 by paleontologists Lü Junchang and Zhang Xingliao. The type species is Eopteranodon lii. A second species, E. yixianensis, was named in 2023, but its validity has been questioned.

<i>Chaoyangopterus</i> Genus of chaoyangopterid pterosaur from the Early Cretaceous

Chaoyangopterus is a genus of chaoyangopterid pterosaur known from a partial skeleton found in Liaoning, China. Chaoyangopterus was found in rocks dating back to the Aptian-age Lower Cretaceous Jiufotang Formation of Dapingfang, Chaoyang.

<i>Aralazhdarcho</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Aralazhdarcho is a genus of azhdarchid pterosaur that lived during the Santonian to the early Campanian ages of the Late Cretaceous period in what is now Kazakhstan. The type and only known species is Aralazhdarcho bostobensis, named and described by Russian paleontologist Alexander Averianov. Its remains were found in the Bostobe Formation of Kazakhstan. The generic name combines the Aral Sea and the related genus Azhdarcho, while the specific name refers to its origin from the Bostobe Formation.

<span class="mw-page-title-main">Azhdarchoidea</span> Superfamily of pterodactyloid pterosaurs

Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. Remains of this group have been found in the Americas, Africa, and Eurasia, suggesting that they probably had a global distribution.

<span class="mw-page-title-main">Chaoyangopteridae</span> Family of azhdarchoid pterosaurs

Chaoyangopteridae is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, Microtuban, Xericeps and Argentinadraco, may extend the fossil range to the Late Cretaceous.

<i>Alanqa</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Alanqa is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Kem Kem Beds of southeastern Morocco. The name Alanqa comes from the Arabic word العنقاءal-‘Anqā’, for a mythical bird of Arabian culture.

<span class="mw-page-title-main">Dsungaripteridae</span> Family of ornithocheiroid pterosaurs

Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings, and were commonly interpreted as durophagous and possibly piscivorous pterosaurs. Fossils have been discovered from Early Cretaceous deposits in Asia, South America and possibly Europe.

<i>Aerotitan</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Aerotitan is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous period in what is now Argentina. Its only remains, which consist of a partial snout, were found in the Allen Formation of the Neuquén Basin in northern Patagonia. This specimen was made the holotype of Aerotitan sudamericanus by paleontologist Fernando Novas and colleagues. The generic name combines the Greek word ἀήρ, meaning "air" and Titan, alluding to the pterosaur's large size. The specific name is a reference to its origin, South America.

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

Banguela is a genus of azhdarchoid pterosaur from the Early Cretaceous period of what is now Brazil. Only one species is known, Banguela oberlii.

<span class="mw-page-title-main">Tapejaroidea</span> Clade of ornithocheiroid pterosaurs

Tapejaroidea is a group of pterosaurs belonging to the clade Ornithocheiroidea. Tapejaroids lived from the Early to Late Cretaceous periods, with one possible member, Tendaguripterus, extending the fossil range to the Late Jurassic period. Tapejaroidea contains two groups, the Dsungaripteridae and the Azhdarchoidea, which in turn includes the azhdarchids, the group that contains some of the largest flying animals. The group was named by Brazilian paleontologist Alexander Wilhelm Armin Kellner in 1996.

<i>Argentinadraco</i> Genus of azhdarchoid pterosaur from the Late Cretaceous

Argentinadraco is an extinct genus of azhdarchoid pterosaur from the Late Cretaceous Portezuelo Formation of Argentina. It contains a single species, A. barrealensis, named in 2017 by Alexander Kellner and Jorge Calvo. Argentinadraco is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the Chaoyangopteridae. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.

<i>Keresdrakon</i> Genus of azhdarchoid pterosaur from the Cretaceous period

Keresdrakon is an extinct genus of azhdarchoid pterosaur from the Goio-Erê Formation of Brazil, which dates back to the Early Cretaceous period, 125 to 100.5 million years ago. Keresdrakon contains a single species, Keresdrakon vilsoni.

<i>Apatorhamphus</i> Genus of azhdarchoid pterosaur from the Late Cretaceous

Apatorhamphus is an extinct genus of azhdarchoid pterosaur from the Kem Kem Group of Morocco. It might have been part of the Chaoyangopteridae. It is only known from a few snout fragments and it likely had a wingspan of between 3–7 metres (9.8–23.0 ft)

<i>Wellnhopterus</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Wellnhopterus is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous period in what us now Texas, United States. Its fossil remains were discovered in the Javelina Formation at Big Bend National Park, located in Brewster County, Texas. The fossils consist of a set of upper and lower jaws, several cervical (neck) vertebrae, and a fragmentary long bone. Originally referred to an indeterminate species of Quetzalcoatlus, these remains would receive various interpretations ranging from a tapejarid to an azhdarchid identity. It was not until December 2021 that they were given a new genus and type species, Wellnhopterus brevirostris, named and described by paleontologists Brian Andres and, posthumously, Wann Langston Jr.. Its generic name means "Wellnhofer's wing", named in honor of paleontologist Peter Wellnhofer, while its specific name means "short-beaked", attributing to the blunt structure of its jaws.

References

  1. ↑ Kellner, A. W. A.; Campos, D. A. (2007). "Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea)". Boletim do Museu Nacional: Geologia. 75: 1–14.
  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. doi:10.1144/GSL.SP.2003.217.01.11. S2CID   86710955.
  3. ↑ Martill, D. M.; Naish, D. (2006). "Cranial crest development in the Azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly". Palaeontology. 49 (4): 925–941. Bibcode:2006Palgy..49..925M. doi: 10.1111/j.1475-4983.2006.00575.x . S2CID   15609202.
  4. ↑ Lü J.; D.M. Unwin; Xu L.; Zhang X. (2008). "A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution". Naturwissenschaften. 95 (9): 891–7. doi:10.1007/s00114-008-0397-5. PMID   18509616.
  5. ↑ Witton, M. P. (2009). "A new species of Tupuxuara (Thalassodromidae, Azhdarchoidea) from the Lower Cretaceous Santana Formation of Brazil, with a note on the nomenclature of Thalassodromidae". Cretaceous Research. 30 (5): 1293–1300. Bibcode:2009CrRes..30.1293W. doi:10.1016/j.cretres.2009.07.006. S2CID   140174098.
  6. ↑ Andres, B.; Clark, J.; Xu, X. (2014). "The Earliest Pterodactyloid and the Origin of the Group". Current Biology . 24 (9): 1011–6. doi: 10.1016/j.cub.2014.03.030 . PMID   24768054.
  7. ↑ Chachere, Vickie (April 24, 2014). "International Scientific Team Discovers, Names Oldest Pterodactyloid Species". USF News. Archived from the original on April 26, 2014. Retrieved April 25, 2014.
  8. 1 2 3 Andres, Brian (December 7, 2021). "Phylogenetic systematics of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea)". Journal of Vertebrate Paleontology. 41 (sup1): 203–217. Bibcode:2021JVPal..41S.203A. doi: 10.1080/02724634.2020.1801703 . ISSN   0272-4634. S2CID   245078533.
  9. 1 2 Longrich, Nicholas R.; Martill, David M.; Andres, Brian; Penny, David (2018). "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary". PLOS Biology. 16 (3): e2001663. doi: 10.1371/journal.pbio.2001663 . PMC   5849296 . PMID   29534059.
  10. ↑ Pinheiro, F.L., Fortier, D.C., Schultz, C.L., De Andrade, J.A.F.G. and Bantim, R.A.M. (in press). "New information on Tupandactylus imperator, with comments on the relationships of Tapejaridae (Pterosauria)." Acta Palaeontologica Polonica, in press, available online 03 Jan 2011. doi : 10.4202/app.2010.0057
  11. ↑ Pêgas, R. V.; Costa, F. R.; Kellner, A. W. A. (2018). "New Information on the osteology and a taxonomic revision of The genus Thalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae)". Journal of Vertebrate Paleontology. 38 (2): e1443273. Bibcode:2018JVPal..38E3273P. doi:10.1080/02724634.2018.1443273. S2CID   90477315.
  12. 1 2 3 4 Pêgas, Rodrigo V. (June 10, 2024). "A taxonomic note on the tapejarid pterosaurs from the Pterosaur Graveyard site (Caiuá Group, ?Early Cretaceous of Southern Brazil): evidence for the presence of two species". Historical Biology : 1–22. doi:10.1080/08912963.2024.2355664. ISSN   0891-2963.
  13. ↑ Cerqueira GM, Santos MA, Marks MF, Sayão JM, Pinheiro FL (2021). "A new azhdarchoid pterosaur from the Lower Cretaceous of Brazil and the paleobiogeography of the Tapejaridae". Acta Palaeontologica Polonica. 66. doi: 10.4202/app.00848.2020 ..
  14. 1 2 Cerqueira, G. M.; Müller, R. T.; Pinheiro, F. L. (2024). "On the phylogenetic affinities of the tapejarid pterosaur 'Tupuxuara deliradamus' from the Lower Cretaceous of Brazil". Historical Biology: An International Journal of Paleobiology. 36 (3): 677–682. doi:10.1080/08912963.2023.2180741.
  15. ↑ Kellner, Alexander W. A.; Weinschütz, Luiz C.; Holgado, Borja; Bantim, Renan A. M.; Sayão, Juliana M. (August 19, 2019). "A new toothless pterosaur (Pterodactyloidea) from Southern Brazil with insights into the paleoecology of a Cretaceous desert". Anais da Academia Brasileira de Ciências. 91 (suppl 2): e20190768. doi: 10.1590/0001-3765201920190768 . ISSN   0001-3765. PMID   31432888.
  16. 1 2 3 4 Ortiz David, Leonardo D.; González Riga, Bernardo J.; Kellner, Alexander W. A. (April 12, 2022). "Thanatosdrakon amaru, gen. ET SP. NOV., a giant azhdarchid pterosaur from the upper Cretaceous of Argentina". Cretaceous Research. 135: 105228. Bibcode:2022CrRes.13705228O. doi:10.1016/j.cretres.2022.105228. S2CID   248140163 . Retrieved April 12, 2022.
  17. ↑ Beccari, Victor; Pinheiro, Felipe Lima; Nunes, Ivan; Anelli, Luiz Eduardo; Mateus, Octávio; Costa, Fabiana Rodrigues (2021). "Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade". PLOS ONE. 16 (8): e0254789. Bibcode:2021PLoSO..1654789B. doi: 10.1371/journal.pone.0254789 . PMC   8386889 . PMID   34432814.
  18. ↑ Bennett, Cristopher S. (2024). "Wing phalanges of a ?thalassodromine pterosaur from the Aptian–Albian Antlers Formation of Texas, USA". Cretaceous Research. 154: 105771. doi:10.1016/j.cretres.2023.105771.
  19. 1 2 Pêgas, R. V.; Zhoi, X.; Jin, X.; Wang, K.; Ma, W. (2023). "A taxonomic revision of the Sinopterus complex (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota, with the new genus Huaxiadraco". PeerJ. 11. e14829. doi: 10.7717/peerj.14829 . PMC   9922500 .
  20. ↑ Zhou, Xuanyu; Ikegami, Naoki; Pêgas, Rodrigo V.; Yoshinaga, Toru; Sato, Takahiro; Mukunoki, Toshifumi; Otani, Jun; Kobayashi, Yoshitsugu (November 16, 2024). "Reassessment of an azhdarchid pterosaur specimen from the Mifune Group, Upper Cretaceous of Japan". Cretaceous Research . 167: 106046. doi:10.1016/j.cretres.2024.106046. ISSN   0195-6671.
  21. 1 2 Pêgas, R.V.; Holgado, B.; Ortiz David, L.D.; Baiano, M.A.; Costa, F.R. (2022). "On the pterosaur Aerotitan sudamericanus (Neuquén Basin, Upper Cretaceous of Argentina), with comments on azhdarchoid phylogeny and jaw anatomy". Cretaceous Research. 129: Article 104998. doi:10.1016/j.cretres.2021.104998. ISSN   0195-6671. S2CID   238725853.
  22. ↑ Smith, Roy E.; Ibrahim, Nizar; Longrich, Nicholas; Unwin, David M.; Jacobs, Megan L.; Williams, Cariad J.; Zouri, Samir; Martill, David M. (2023). "The pterosaurs of the Cretaceous Kem Kem Group of Morocco". PalZ. 97: 519–568. doi:10.1007/s12542-022-00642-6.
  23. ↑ Pêgas, R. V.; Zhoi, X.; Jin, X.; Wang, K.; Ma, W. (2023). "A taxonomic revision of the Sinopterus complex (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota, with the new genus Huaxiadraco". PeerJ. 11. e14829. doi: 10.7717/peerj.14829 . PMC   9922500 .
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