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Thalassodromidae

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Thalassodromids
Temporal range: Early-Late Cretaceous, 112–88 Ma
Replica skeleton of Tupuxuara leonardii
Scientific classification Edit this classification
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

[edit]
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

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]

References

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  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. ^ a b c 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. ^ a b 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. ^ a b c d 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. ^ a b 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. ^ a b c d 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. ^ a b 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. ^ a b 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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