Dsungaripteridae

Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea.^[2] They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings,^[3] and were commonly interpreted as durophagous and possibly piscivorous pterosaurs.^[4] Fossils have been discovered from Early Cretaceous deposits in Asia, South America and possibly Europe.^[5]

Dsungaripterids Temporal range: Early Cretaceous PreꞒ Ꞓ O S D C P T J K Pg N
Restored skeleton of Dsungaripterus weii
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Order:	†Pterosauria
Suborder:	†Pterodactyloidea
Clade:	†Ornithocheiroidea
Family:	†Dsungaripteridae Young, 1964
Type species
†Dsungaripterus weii Young, 1964
Genera
†Lonchognathosaurus †Noripterus †Ordosipterus †Puntanipterus?[1] †Dsungaripterinae †Domeykodactylus †Dsungaripterus

Classification

In 1964, Young created a family to place the genus Dsungaripterus,^[6] a Chinese taxon with potential remains also known from the Hasandong Formation of South Korea.^[7][8] Later on, Noripterus (then now with the name "Phobetor" which was already occupied, therefore the quotation marks) was also assigned to the family.^[9]

In 2003, Alexander Kellner gave the exact definition as a clade:^[10] the group was composed out of the latest common ancestor of Dsungaripterus, Noripterus and “Phobetor”, and all its descendants. As synapomorphies he gave the next six characteristics: a relatively small eye-socket, which is placed high up the skull; an opening below the eye-socket; a high ridge across the snout, which starts in front of the nasal opening and ends behind the eye-sockets; the maxilla reaches out down- and backwards; the absence of teeth in the first part of the jaws; the teeth in the back of the upper jaw are the biggest; the teeth have a wide oval basis. Kellner pointed out all members of the group, except for Dsungaripterus itself, were known from fragmentary remains, so only the last characteristic could be established for sure in all members.^[10]

In the same year, David Unwin gave a slightly different definition: the last common ancestor of Dsungaripterus weii and Noripterus complicidens, and all its descendants.^[11] The group belongs to the Dsungaripteroidea sensu Unwin and is presumably relatively closely related to the Azhdarchoidea. According to Unwin, Germanodactylus is the sister taxon to the group, but his analyses have this outcome as the only ones. According to an analysis by Brian Andres from 2008, the Dsungaripteridae are closely related to the Tapejaridae, what would actually make them members of the Azhdarchoidea.^[12] Recent examinations of the palate of Dsungaripterus support an azhdarchoid interpretation.^[13]

The earliest possible fossils attributed to this group are from the Berriasian age of Europe based on two specimens found in Romania, MTCO 17.764/2019 and MTCO 18.262/9651,^[5] though the latter specimen has been reinterpreted as an indeterminate azhdarchoid.^[14] Previous Late Jurassic records have been disputed,^[15] and a possible dsungaripterid specimen from the Kimmeridgian of Germany has been suggested as an indeterminate dsungaripteroid or a specimen of Germanodactylus.^[16][17] The earliest known taxon is from the Early Cretaceous of Chile, belonging to the species Domeykodactylus ceciliae.^[15] Among the latest known dsungaripterid species is Lonchognathosaurus acutirostris, from the Albian-age Lower Cretaceous Lianmuqin Formation of Xinjiang, China, about 112 million years ago.^[18]

Family tree

Below is a cladogram showing the results of a phylogenetic analysis presented by Andres and colleagues in 2014. They recovered Dsungaripteridae within the Dsungaripteromorpha (a subgroup within the Azhdarchoidea), sister taxon to the subfamily Thalassodrominae. The found Dsungaripteridae to consist of two subfamilies called Dsungaripterinae and Noripterinae (which consists only of Noripterus species). Their cladogram is shown below.^[19]

Dsungaripteromorpha	Dsungaripteridae Dsungaripterinae Dsungaripterus weii Domeykodactylus ceciliae Noripterinae Noripterus parvus Noripterus complicidens Thalassodrominae Thalassodromeus sethi Tupuxuara longicristatus Tupuxuara leonardii

In 2019, a different topology, this time by Kellner and colleagues, was published. In this study, Dsungaripteridae was recovered outside the Azhdarchoidea, within the larger group Tapejaroidea. The cladogram of the analysis is shown below.^[20]

Tapejaroidea	Dsungaripteridae Dsungaripterus weii Noripterus parvus Azhdarchoidea Azhdarchidae Chaoyangopteridae Tapejaromorpha Keresdrakon vilsoni Tapejaridae Thalassodrominae Tapejarinae

References

1. Bonaparte, J.F., and Sanchez, T.M. (1975). Restos de un pterosaurio Puntanipterus globosus de la formación La Cruz provincia San Luis, Argentina. Actas Primo Congresso Argentino de Paleontologia e Biostratigraphica 2:105-113. [Spanish]
2. Unwin, David M. (2006). The Pterosaurs: From Deep Time. New York: Pi Press. p. 273. ISBN 0-13-146308-X.
3. Witton, Mark (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. p. 51. ISBN 978-0691150611.
4. Bestwick, Jordan; Unwin, David M.; Butler, Richard J.; Henderson, Donald M.; Purnell, Mark A. (November 2018). "Pterosaur dietary hypotheses: a review of ideas and approaches". Biological Reviews. 93 (4): 2021–2048. doi:10.1111/brv.12431. PMC 6849529. PMID 29877021.
5. Dyke, G.; Benton, M.; Posmosanu, E.; Naish, D. (2010). "Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania". Palaeontology. 54: 79–95. doi:10.1111/j.1475-4983.2010.00997.x. S2CID 15172374.
6. Young, C.-C. (1964). "On a new pterosaurian from Sinkiang, China" (PDF). Vertebrate PalAsiatica: 221–225.
7. Yang, S.Y. (2015). "Study history and research ethics of the dinosaur, pterosaur and bird tracks from Korea". Journal of the Geological Society of Korea. 51 (2): 127–140. doi:10.14770/jgsk.2015.51.2.127.
8. Kim, Jeong Yul; Huh, Min (2018). Dinosaurs, Birds, and Pterosaurs of Korea: A Paradise of Mesozoic Vertebrates (PDF). Springer Nature. ISBN 978-981-10-6998-7.
9. Lü, J.; Azuma, Y.; Dong, Z.; Barsbold, R.; Kobayashi, Y.; Lee, Y.-N. (2009). "New material of dsungaripterid pterosaurs (Pterosauria: Pterodactyloidea) from western Mongolia and its palaeoecological implications". Geological Magazine. 146 (5): 690–700. doi:10.1017/S0016756809006414.
10. Kellner, A.W.A., 2003. Pterosaur phylogeny and comments on the evolutionary history of the AN group. In: Buffetaut, E., Mazin, J.M. (Eds.), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publication 217, 105–137.
11. Unwin, D. M., (2003). "On the phylogeny and evolutionary history of pterosaurs." Pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347.
12. Andres, B.; Ji, Q. (2008). "A new pterosaur from the Liaoning Province of China, the phylogeny of the Pterodactyloidea, and convergence in their cervical vertebrae". Palaeontology. 51 (2): 453–469. doi:10.1111/j.1475-4983.2008.00761.x.
13. Chen, He; Jiang, Shunxing; Kellner, Alexander W. A.; Cheng, Xin; Zhang, Xinjun; Qiu, Rui; Li, Yang; Wang, Xiaolin (April 1, 2020). "New anatomical information on Dsungaripterus weii Young, 1964 with focus on the palatal region". PeerJ. 8: e8741. doi:10.7717/peerj.8741. PMC 7127482. PMID 32274262.
14. Alexander Averianov (2014). "Review of taxonomy, geographic distribution, and paleoenvironments of Azhdarchidae (Pterosauria)". ZooKeys (432): 1–107. doi:10.3897/zookeys.432.7913. PMC 4141157. PMID 25152671.
15. Martill, D.M.; Frey, E.; Diaz, G.C.; Bell, C.M. (2000). "Reinterpretation of a Chilean pterosaur and the occurrence of Dsungeripteridae in South America" (PDF). Geological Magazine. 137 (1): 19–25. Bibcode:2000GeoM..137...19M. doi:10.1017/S0016756800003502. S2CID 128593191.
16. Hyder, E. S.; Witton, M. P.; Martill, D. M. (2014). "Evolution of the Pterosaur Pelvis". Acta Palaeontologica Polonica. 59 (1): 109–124. doi:10.4202/app.2011.1109. S2CID 55405758.
17. Alexander O. Averianov; Stepan V. Ivantsov; Sergey V. Leshchinskiy; Pavel P.Skutschas (12 April 2022). "First pterosaur bone from the Lower Cretaceous of Siberia, Russia". Cretaceous Research. 135: 105230. doi:10.1016/j.cretres.2022.105230. S2CID 248136173. Retrieved 13 April 2022.
18. Maisch, M.W.; Matzke, A.T.; Sun, Ge (2004). "A new dsungaripteroid pterosaur from the Lower Cretaceous of the southern Junggar Basin, north-west China" (PDF). Cretaceous Research. 25 (5): 625–634. doi:10.1016/j.cretres.2004.06.002.
19. 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.
20. Kellner, Alexander W. A.; Weinschütz, Luiz C.; Holgado, Borja; Bantim, Renan A. M.; Sayão, Juliana M. (19 August 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.