thalassodromidae

File:Kariridraco_by_Júlia_d’Oliveira.jpg

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.{{cite journal |last1=Kellner |first1=A. W. A. |last2=Campos |first2=D. A. |title=Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea) |journal=Boletim do Museu Nacional: Geologia |volume=75 |pages=1–14 |year=2007}}

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.{{cite journal |last1=Unwin |first1=D. M. |title=On the phylogeny and evolutionary history of pterosaurs |journal=Geological Society, London, Special Publications |date=2003 |volume=217 |issue=1 |pages=139–190 |doi=10.1144/GSL.SP.2003.217.01.11 |bibcode=2003GSLSP.217..139U |s2cid=86710955}} 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.{{cite journal |last1=Martill |first1=D. M. |last2=Naish |first2=D. |title=Cranial crest development in the Azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly |journal=Palaeontology |date=2006 |volume=49 |issue=4 |pages=925–941 |doi=10.1111/j.1475-4983.2006.00575.x |s2cid=15609202 |doi-access=free |bibcode=2006Palgy..49..925M}} 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.{{cite journal |author=Lü J. |author2=D.M. Unwin |author3=Xu L. |author4=Zhang X. |year=2008 |title=A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution |journal=Naturwissenschaften |pages=891–7 |doi=10.1007/s00114-008-0397-5 |volume=95 |pmid=18509616 |issue=9}} 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.{{cite journal |last1=Witton |first1=M. P. |title=A new species of Tupuxuara (Thalassodromidae, Azhdarchoidea) from the Lower Cretaceous Santana Formation of Brazil, with a note on the nomenclature of Thalassodromidae |journal=Cretaceous Research |date=2009 |volume=30 |issue=5 |pages=1293–1300 |doi=10.1016/j.cretres.2009.07.006|bibcode=2009CrRes..30.1293W |s2cid=140174098 }} 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.{{cite journal |author1=Andres, B. |author2=Clark, J. |author3=Xu, X. |year=2014 |title=The Earliest Pterodactyloid and the Origin of the Group |journal=Current Biology |volume= 24|issue=9 |pages= 1011–6|url=https://www.cell.com/current-biology/pdfExtended/S0960-9822(14)00322-4|doi=10.1016/j.cub.2014.03.030 |pmid=24768054|doi-access=free }}{{cite web |url=http://news.usf.edu/article/templates/?a=6251&z=123 |title=International Scientific Team Discovers, Names Oldest Pterodactyloid Species |first=Vickie |last=Chachere |date=April 24, 2014 |work=USF News |access-date=April 25, 2014 |archive-date=April 26, 2014 |archive-url=https://web.archive.org/web/20140426232825/http://news.usf.edu/article/templates/?a=6251&z=123 |url-status=dead }}

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.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}} Studies by Rodrigo Pêgas and colleagues,{{cite journal |last1=Pêgas |first1=R. V. |last2=Costa |first2=F. R. |last3=Kellner |first3=A. W. A. |title=New Information on the osteology and a taxonomic revision of The genus Thalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae) |journal=Journal of Vertebrate Paleontology |date=2018 |volume=38 |issue=2 |pages=e1443273 |doi=10.1080/02724634.2018.1443273 |bibcode=2018JVPal..38E3273P |s2cid=90477315}} Gabriela Cerquiera and colleagues,{{cite journal|vauthors=Cerqueira GM, Santos MA, Marks MF, Sayão JM, Pinheiro FL|title=A new azhdarchoid pterosaur from the Lower Cretaceous of Brazil and the paleobiogeography of the Tapejaridae|year=2021|journal=Acta Palaeontologica Polonica|volume=66|doi=10.4202/app.00848.2020|doi-access=free|url=https://www.app.pan.pl/article/item/article/item/app008482020.html?pdf=39}}. as well as Kellner himself have continued to followed Kellner's original model, into the 2020s,{{Cite journal |last1=Kellner |first1=Alexander W. A. |last2=Weinschütz |first2=Luiz C. |last3=Holgado |first3=Borja |last4=Bantim |first4=Renan A. M. |last5=Sayão |first5=Juliana M. |date=August 19, 2019 |title=A new toothless pterosaur (Pterodactyloidea) from Southern Brazil with insights into the paleoecology of a Cretaceous desert |journal=Anais da Academia Brasileira de Ciências |language=en |volume=91 |issue=suppl 2 |pages=e20190768 |doi=10.1590/0001-3765201920190768 |pmid=31432888 |issn=0001-3765 |doi-access=free}} amongst others.{{cite journal |last1=Beccari |first1=Victor |last2=Pinheiro |first2=Felipe Lima |last3=Nunes |first3=Ivan |last4=Anelli |first4=Luiz Eduardo |last5=Mateus |first5=Octávio |last6=Costa |first6=Fabiana Rodrigues |title=Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade |journal=PLOS ONE |date=2021 |volume=16 |issue=8 |pages=e0254789 |pmid=34432814|doi=10.1371/journal.pone.0254789|pmc=8386889 |bibcode=2021PLoSO..1654789B |doi-access=free }}{{cite journal | url=https://www.sciencedirect.com/science/article/pii/S0195667123002999?casa_token=JLd2zJhI1I0AAAAA:-OpizuH51C1utc7lpoG4bnI3WDslPzh3-KYtFHgHrdFKwdjnYuHs1MlNlIWpG5qp_wtK6BQ5zcPn | title=Wing phalanges of a ?thalassodromine pterosaur from the Aptian–Albian Antlers Formation of Texas, USA | author=Bennett, Cristopher S. | journal=Cretaceous Research | year=2024 | volume=154 | pages=105771 | doi=10.1016/j.cretres.2023.105771}} 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.{{Cite journal |last1=Pêgas |first1=R. V. |last2=Zhoi |first2=X. |last3=Jin |first3=X. |last4=Wang |first4=K. |last5=Ma |first5=W. |year=2023 |title=A taxonomic revision of the Sinopterus complex (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota, with the new genus Huaxiadraco |journal=PeerJ |volume=11 |at=e14829 |doi=10.7717/peerj.14829 |doi-access=free |pmc=9922500}} Subsequent studies by Pêgas and colleagues would follow this change,{{Cite journal |last1=Pêgas |first1=Rodrigo V. |date=June 10, 2024 |title=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 |journal=Historical Biology |language=en |pages=1–22 |url=https://www.researchgate.net/publication/381306918 |doi=10.1080/08912963.2024.2355664 |issn=0891-2963}}{{Cite journal |last1=Zhou |first1=Xuanyu |last2=Ikegami |first2=Naoki |last3=Pêgas |first3=Rodrigo V. |last4=Yoshinaga |first4=Toru |last5=Sato |first5=Takahiro |last6=Mukunoki |first6=Toshifumi |last7=Otani |first7=Jun |last8=Kobayashi |first8=Yoshitsugu |date=November 16, 2024 |title=Reassessment of an azhdarchid pterosaur specimen from the Mifune Group, Upper Cretaceous of Japan |url=https://linkinghub.elsevier.com/retrieve/pii/S0195667124002192 |journal=Cretaceous Research |volume=167 |pages=106046 |doi=10.1016/j.cretres.2024.106046 |issn=0195-6671}} though others continue to use the traditional scope of Tapejaridae.{{Cite journal |last1=Cerqueira |first1=G. M. |last2=Müller |first2=R. T. |last3=Pinheiro |first3=F. L. |year=2024 |title=On the phylogenetic affinities of the tapejarid pterosaur {{‘}}Tupuxuara deliradamus{{’}} from the Lower Cretaceous of Brazil |journal=Historical Biology: An International Journal of Paleobiology |volume=36 |issue=3 |pages=677–682 |doi=10.1080/08912963.2023.2180741}}

{{col-begin|width=95%}}

{{col-2}}

Unwin-model cladogram based on Andres, 2021:

{{clade|style=font-size:85%;line-height:85%

|label1=Azhdarchoidea

|1={{clade

|1=Tapejaridae

|label2=Neoazhdarchia

|2={{clade

|1=Neopterodactyloidea

|label2=Dsungaripteromorpha

|2={{clade

|1=Microtuban altivolans

|2={{clade

|1=Dsungaripteridae

|label2=Thalassodromidae

|2={{clade

|1={{clade

|1=Tupuxuara longicristatus

|2=Tupuxuara leonardii

}}

|label2=Thalassodrominae

|2={{clade

|1=Thalassodromeus sethi

|2={{clade

|1=Alanqa saharica

|2={{clade

|1=Argentinadraco barrealensis

|2={{clade

|1=Leptostomia begaaensis

|2=Xericeps curvirostris

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

{{col-2}}

Kellner-model cladogram based on Pêgas and colleagues, 2023:

{{clade|style=font-size:85%;line-height:85%

|label1=Tapejaroidea

|1={{clade

|1=Dsungaripteridae

|label2=Azhdarchoidea

|2={{clade

|1=Azhdarchomorpha

|label2=Tapejaromorpha

|2={{clade

|label1=Tapejaridae

|1={{clade

|1=Caupedactylia

|2=Eutapejaria

}}

|label2=Thalassodromidae

|2={{clade

|1={{clade

|1=Thalassodromeus sethi

|2=Thalassodromeus oberlii

}}

|2={{clade

|1=Kariridraco dianae

|2={{clade

|1=Tupuxuara longicristatus

|2=Tupuxuara leonardii

}} }} }} }} }} }} }}

{{col-end}}

File:Thalassodromeus_skull.png; 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.{{cite journal |last1=Longrich |first1=Nicholas R. |last2=Martill |first2=David M. |last3=Andres |first3=Brian |last4=Penny |first4=David |title=Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary |journal=PLOS Biology |date=2018 |volume=16 |issue=3 |pages=e2001663 |doi=10.1371/journal.pbio.2001663 |pmid=29534059 |pmc=5849296 |doi-access=free}} 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.{{Cite journal |last=Andres |first=Brian |date=December 7, 2021 |title=Phylogenetic systematics of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea) |journal=Journal of Vertebrate Paleontology |volume=41 |issue=sup1 |pages=203–217 |doi=10.1080/02724634.2020.1801703 |s2cid=245078533 |issn=0272-4634 |doi-access=free |bibcode=2021JVPal..41S.203A}} Other studies would not corroborate these results. In a 2022 study, Pêgas and colleagues re-evaluated Aerotitan and rejected its identity as a thalassodromid. Other studies have supported this conclusion.{{cite journal |last1=Ortiz David |first1=Leonardo D. |last2=González Riga |first2=Bernardo J. |last3=Kellner |first3=Alexander W. A. |date=April 12, 2022 |title=Thanatosdrakon amaru, gen. ET SP. NOV., a giant azhdarchid pterosaur from the upper Cretaceous of Argentina |url=https://www.sciencedirect.com/science/article/abs/pii/S0195667122000921 |journal=Cretaceous Research |volume=135 |page=105228 |doi=10.1016/j.cretres.2022.105228 |bibcode=2022CrRes.13705228O |s2cid=248140163 |access-date=April 12, 2022}} 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.{{Cite journal |last1=Pêgas |first1=R.V. |last2=Holgado |first2=B. |last3=Ortiz David |first3=L.D. |last4=Baiano |first4=M.A. |last5=Costa |first5=F.R. |url=https://www.sciencedirect.com/science/article/abs/pii/S0195667121002469 |title=On the pterosaur Aerotitan sudamericanus (Neuquén Basin, Upper Cretaceous of Argentina), with comments on azhdarchoid phylogeny and jaw anatomy |year=2022 |journal=Cretaceous Research |volume=129 |pages=Article 104998 |doi=10.1016/j.cretres.2021.104998 |issn=0195-6671 |s2cid=238725853}} 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.{{cite journal | url=https://link.springer.com/article/10.1007/s12542-022-00642-6 | title=The pterosaurs of the Cretaceous Kem Kem Group of Morocco | journal=PalZ | last1=Smith | first1=Roy E. | last2=Ibrahim | first2=Nizar | last3=Longrich | first3=Nicholas | last4=Unwin | first4=David M. | last5=Jacobs | first5=Megan L. | last6=Williams | first6=Cariad J. | last7=Zouri | first7=Samir | last8=Martill | first8=David M. | year=2023 | volume=97 | pages=519–568 | doi=10.1007/s12542-022-00642-6| doi-access=free }} Subsequent studies have placed these two genera and Argentinadraco in Alanqidae, Azhdarchidae, or Chaoyangopteridae.{{Cite journal|last1=Pêgas |first1=R. V. |last2=Zhoi |first2=X. |last3=Jin |first3=X. |last4=Wang |first4=K. |last5=Ma |first5=W. |year=2023 |title=A taxonomic revision of the Sinopterus complex (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota, with the new genus Huaxiadraco |journal=PeerJ |volume=11 |at=e14829 |doi=10.7717/peerj.14829 |doi-access=free |pmc=9922500 }}

{{Reflist}}

{{Pterosauria|Az.}}

{{Taxonbar|from1=Q7709559|from2=Q117038917}}

Category:Azhdarchoidea

Category:Albian first appearances

Category:Maastrichtian extinctions