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Tsogtopteryx

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Tsogtopteryx
Holotype cervical vertebra VI
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: †Pterosauria
Suborder: †Pterodactyloidea
Clade: †Azhdarchoidea
Family: †Azhdarchidae
Clade: †Hatzegopterygia
Genus: †Tsogtopteryx
Pêgas, Zhou & Kobayashi, 2025
Species:
†T. mongoliensis
Binomial name
†Tsogtopteryx mongoliensis
Pêgas, Zhou & Kobayashi, 2025

Tsogtopteryx (lit. 'mighty hero wing') is an extinct genus of azhdarchid pterosaurs known from the Late Cretaceous Bayanshiree Formation of Mongolia. The genus contains a single species, Tsogtopteryx mongoliensis, known from most of a cervical (neck) vertebra. It was closely related to Hatzegopteryx, and coexisted with Gobiazhdarcho, another azhdarchid more closely related to Quetzalcoatlus.

Discovery and naming

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Mongolian Cretaceous fossil localities; Tsogtopteryx is known from the Bayshin Tsav locality in Area C (Bayanshiree Formation)

In 1993, a joint paleontological expedition between the Hayashibara Museum of Natural Sciences and the Mongolian Paleontological Center conducted fieldwork in the Gobi Desert of Mongolia. At the 'Bayshin Tsav' locality of the Bayanshiree Formation in Dornogovi Province, workers collected an isolated cervical (neck) vertebral element. This specimen, accessioned as MPC−Nd 100/303, comprises almost all of the sixth cervical vertebra, missing the posteriormost (furthest back) region.[1] In 2009, Mahito Watabe and colleagues described this specimen, in addition to another isolated pterosaur cervical vertebra from the 'Burkhant' locality of the same formation. The authors refrained from naming either specimen, but discussed their anatomy and likely phylogenetic affinities in depth.[2]

In 2025, R. V. Pêgas, Xuanyu Zhou, and Yoshitsugu Kobayashi described Tsogtopteryx mongoliensis as a new genus and species of pterosaurs based on the 'Bayshin Tsav azhdarchid' fossil remains. The generic name, Tsogtopteryx, combines the Mongolian tsogt, meaning "mighty hero", with the Ancient Greek pteryx, meaning 'wing'. The specific name, mongoliensis, references the discovery of the specimen in Mongolia. In the same publication, the authors also named the Burkhant specimen as another new azhdarchid, Gobiazhdarcho.[1]

Prior to the naming of Tsogtopteryx and Gobiazhdarcho, the only pterosaur named from Mongolia was the dsungaripterid Noripterus parvus,[3] known from the Tsagan-Tsab Formation.[1] Several fragmentary indeterminate pterosaur remains are also known from the country, including a possible anurognathid, anhanguerid,[4] and tapejaroid,[5] in addition to a possible azhdarchid long bone found in the stomach of a dromaeosaurid dinosaur and the very fragmentary remains of a giant azhdarchid.[6][7]

Description

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Based on the size of the holotype vertebra (more than 7.7 centimetres (3.0 in) long), the wingspan of Tsogtopteryx was estimated at 1.6–1.9 metres (5.2–6.2 ft), which is remarkably small for an azhdarchid. This would make it the smallest named azhdarchid, only smaller than an unnamed possible member of the clade from the Northumberland Formation of Canada, which has an estimated wingspan of 1.6 m (5.2 ft) or less.[8] Despite this diminutive size, the specimen likely belongs to a skeletally mature (fully grown) individual. This was determined based on the lack of a grained texture often seen in immature pterosaur bones and the full fusion of the cervical ribs to the vertebra, which does not happen until maturity. In comparison, the coeval Gobiazhdarcho was not yet fully grown but was notably larger with a 3–3.5 metres (9.8–11.5 ft) wingspan.[1]

Classification

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Life restoration of the related Cryodrakon

To determine the relationships and affinities of Tsogtopteryx, Pêgas and colleagues scored it in a comprehensive pterosaur-focused phylogenetic matrix modified from Zhou et al. (2025),[9] deriving from Pêgas (2024)[3] and other earlier publications, with modifications based on newer literature. Their analyses recovered Tsogtopteryx as the basalmost member of the clade Hatzegopterygia, which includes azhdarchids more closely related to Hatzegopteryx than Quetzalcoatlus. The results are displayed in the cladogram below:[1]

Azhdarchidae

Paleoenvironment

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Speculative life restoration of Tsogtopteryx (smaller; blue) and Gobiazhdarcho (larger; red) in a Bayanshiree environment with the ornithopod dinosaur Gobihadros

Tsogtopteryx is known from the 'Bayshin Tsav' locality of the Bayanshiree Formation. Examinations of the magnetostratigraphy of the formation confirm that it lies entirely within the Cretaceous Long Normal, which lasted only until the end of the Santonian stage. Calcite U–Pb measurements estimate the age of the Bayanshiree Formation from 95.9 ± 6.0 million to 89.6 ± 4.0 million years ago, in the Albian through Santonian ages.[12][13] Considering all available evidence, there are likely two distinct levels of the formation—an 'upper' and 'lower'—partially based on faunal differences: a lower part lasting from the Cenomanian to late Turonian ages and an upper part lasting the late Turonian to Santonian ages in the late Cretaceous period.[14] The Bayshin Tsav locality is part of the upper Bayanshiree.[1]

Fluvial, lacustrine and caliche-based sedimentation indicates a lesser semi-arid climate, with the presence of wet environments composed of large meanders and lakes. Large-scale cross-stratification in many of the sandstone layers at the Baynshire and Burkhant localities seems to indicate large meandering rivers, and these large water bodies may have drained the eastern part of the Gobi Desert.[15][12]

A vast faunal diversity is known in the formation, comprising dinosaur and non-dinosaur genera. Dinosaurs definitively recognized from the Bayshin Tsav locality include the tyrannosauroid Khankhuuluu,[16] the ornithomimosaur Garudimimus,[17] the therizinosaurs Erlikosaurus and Segnosaurus, the ankylosaur Talarurus, and the hadrosauroid Gobihadros.[18][1] Other theropod dinosaurs from other localities include the additional therizinosaurs Duonychus and Enigmosaurus,[19] and the large dromaeosaurid Achillobator.[20] Herbivorous dinosaurs from other localities are represented by the ankylosaur Tsagantegia,[21] the small marginocephalians Amtocephale (a pachycephalosaur) and Graciliceratops (a ceratopsian),[22][23] and the sauropod Erketu.[24] Other fauna include semiaquatic reptiles like crocodylomorphs and nanhsiungchelyid turtles, and various fish.[14][25] Numerous fossilized fruits have been recovered from the Bor Guvé and Khara Khutul localities.[24]

References

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  1. ^ a b c d e f g Pêgas, R. V.; Zhou, Xuanyu; Kobayashi, Yoshitsugu (2025). "Azhdarchid pterosaur diversity in the Bayanshiree Formation, Upper Cretaceous of the Gobi Desert, Mongolia". PeerJ. 13 e19711. doi:10.7717/peerj.19711.
  2. ^ Watabe, Mahito; Tsuihiji, Takanobu; Suzuki, Shigeru; Tsogtbaatar, Khishigjav (2009). "The First Discovery of Pterosaurs from the Upper Cretaceous of Mongolia" (PDF). Acta Palaeontologica Polonica. 54 (2): 231–242. doi:10.4202/app.2006.0068. ISSN 0567-7920.
  3. ^ a b Pêgas, Rodrigo V. (2024-06-10). "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.
  4. ^ Bakhurina, Natalia N.; Unwin, David M. (1995). "A survey of pterosaurs from the Jurassic and Cretaceous of the former soviet union and Mongolia". Historical Biology. 10 (3): 197–245. doi:10.1080/10292389509380522. ISSN 0891-2963.
  5. ^ Andres, Brian; Norell, Mark A. (2005-03-24). "The First Record of a Pterosaur from the Early Cretaceous Strata of Öösh (Övörkhangai; Mongolia)". American Museum Novitates. 3472: 1–6. doi:10.1206/0003-0082(2005)472<0001:TFROAP>2.0.CO;2. ISSN 0003-0082.
  6. ^ Hone, David; Tsuihiji, Takanobu; Watabe, Mahito; Tsogtbaatr, Khishigjaw (2012-05-01). "Pterosaurs as a food source for small dromaeosaurs". Palaeogeography, Palaeoclimatology, Palaeoecology. 331–332: 27–30. doi:10.1016/j.palaeo.2012.02.021.
  7. ^ Tsuihiji, Takanobu; Andres, Brian; O'connor, Patrick M.; Watabe, Mahito; Tsogtbaatar, Khishigjav; Mainbayar, Buuvei (September 2017). "Gigantic pterosaurian remains from the Upper Cretaceous of Mongolia". Journal of Vertebrate Paleontology. 37 (5) e1361431. doi:10.1080/02724634.2017.1361431. ISSN 0272-4634.
  8. ^ Martin-Silverstone, Elizabeth; Witton, Mark P.; Arbour, Victoria M.; Currie, Philip J. (August 2016). "A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants". Royal Society Open Science. 3 (8) 160333. doi:10.1098/rsos.160333. ISSN 2054-5703. PMC 5108964. PMID 27853614.
  9. ^ Zhou, Xuanyu; Ikegami, Naoki; Pêgas, Rodrigo V.; Yoshinaga, Toru; Sato, Takahiro; Mukunoki, Toshifumi; Otani, Jun; Kobayashi, Yoshitsugu (March 2025). "Reassessment of an azhdarchid pterosaur specimen from the Mifune Group, Upper Cretaceous of Japan". Cretaceous Research. 167 106046. Bibcode:2025CrRes.16706046Z. doi:10.1016/j.cretres.2024.106046. ISSN 0195-6671.
  10. ^ Vremir, Mátyás; Witton, Mark; Naish, Darren; Dyke, Gareth; Brusatte, Stephen L.; Norell, Mark; Totoianu, Radu (March 17, 2015). "A Medium-Sized Robust-Necked Azhdarchid Pterosaur (Pterodactyloidea: Azhdarchidae) from the Maastrichtian of Pui (Haţeg Basin, Transylvania, Romania)". American Museum Novitates (3827): 1–16. doi:10.1206/3827.1. ISSN 0003-0082.
  11. ^ 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.
  12. ^ a b Hicks, J. F.; Brinkman, D. L.; Nichols, D. J.; Watabe, M. (1999). "Paleomagnetic and palynologic analyses of Albian to Santonian strata at Bayn Shireh, Burkhant, and Khuren Dukh, eastern Gobi Desert, Mongolia". Cretaceous Research. 20 (6): 829–850. Bibcode:1999CrRes..20..829H. doi:10.1006/cres.1999.0188. Archived from the original on 2022-05-06. Retrieved 2020-01-26.
  13. ^ Kurumada, Y.; Aoki, S.; Aoki, K.; Kato, D.; Saneyoshi, M.; Tsogtbaatar, K.; Windley, B. F.; Ishigaki, S. (2020). "Calcite U–Pb age of the Cretaceous vertebrate-bearing Bayn Shire Formation in the Eastern Gobi Desert of Mongolia: usefulness of caliche for age determination". Terra Nova. 32 (4): 246–252. Bibcode:2020TeNov..32..246K. doi:10.1111/ter.12456.
  14. ^ a b Averianov, Alexander; Sues, Hans-Dieter (April 2012). "Correlation of Late Cretaceous continental vertebrate assemblages in Middle and Central Asia". Journal of Stratigraphy. 36 (2): 462–485. hdl:10088/19377.
  15. ^ Samoilov, V. S.; Benjamini, C. (1996). "Geochemical features of dinosaur remains from the Gobi Desert, South Mongolia". PALAIOS. 11 (6): 519–531. Bibcode:1996Palai..11..519S. doi:10.2307/3515188. JSTOR 3515188.
  16. ^ Voris, Jared T.; Zelenitsky, Darla K.; Kobayashi, Yoshitsugu; Modesto, Sean P.; Therrien, François; Tsutsumi, Hiroki; Chinzorig, Tsogtbaatar; Tsogtbaatar, Khishigjav (2025-06-11). "A new Mongolian tyrannosauroid and the evolution of Eutyrannosauria". Nature. doi:10.1038/s41586-025-08964-6.
  17. ^ Kobayashi, Y.; Barsbold, R. (2005). "Reexamination of a primitive ornithomimosaur, Garudimimus brevipes Barsbold, 1981 (Dinosauria: Theropoda), from the Late Cretaceous of Mongolia". Canadian Journal of Earth Sciences. 42 (9): 1501–1521. Bibcode:2005CaJES..42.1501K. doi:10.1139/e05-044. hdl:2115/14579.
  18. ^ Tsogtbaatar, K.; Weishampel, D. B.; Evans, D. C.; Watabe, M. (2019). "A new hadrosauroid (Dinosauria: Ornithopoda) from the Late Cretaceous Baynshire Formation of the Gobi Desert (Mongolia)". PLOS ONE. 14 (4) e0208480. Bibcode:2019PLoSO..1408480T. doi:10.1371/journal.pone.0208480. PMC 6469754. PMID 30995236.
  19. ^ Kobayashi, Yoshitsugu; Zelenitsky, Darla K.; Fiorillo, Anthony R.; Chinzorig, Tsogtbaatar (2025-03-25). "Didactyl therizinosaur with a preserved keratinous claw from the Late Cretaceous of Mongolia". iScience. 28 (4). Bibcode:2025iSci...28k2141K. doi:10.1016/j.isci.2025.112141. ISSN 2589-0042. PMC 12131255.
  20. ^ Perle, A.; Norell, M. A.; Clark, J. (1999). "A new maniraptoran Theropod – Achillobator giganticus (Dromaeosauridae) – from the Upper Cretaceous of Burkhant, Mongolia". Contributions from the Geology and Mineralogy Chair, National Museum of Mongolia (101): 1–105. OCLC 69865262.
  21. ^ Park, J. (2020). "Additional skulls of Talarurus plicatospineus (Dinosauria: Ankylosauridae) and implications for paleobiogeography and paleoecology of armored dinosaurs". Cretaceous Research. 108 e104340. Bibcode:2020CrRes.10804340P. doi:10.1016/j.cretres.2019.104340. S2CID 212423361.
  22. ^ Sereno, P. C. (2000). "The fossil record, systematics and evolution of pachycephalosaurs and ceratopsians from Asia" (PDF). The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press. pp. 489–491. Archived (PDF) from the original on 2019-12-25. Retrieved 2020-01-26.
  23. ^ Watabe, M.; Tsogtbaatar, K.; Sullivan, R. M. (2011). "A new pachycephalosaurid from the Baynshire Formation (Cenomanian-late Santonian), Gobi Desert, Mongolia" (PDF). Fossil Record 3. New Mexico Museum of Natural History and Science, Bulletin. 53: 489–497. Archived (PDF) from the original on 2015-09-24. Retrieved 2020-01-26.
  24. ^ a b Ksepka, D. T.; Norell, M. A. (2006). "Erketu ellisoni, a long-necked sauropod from Bor Guvé (Dornogov Aimag, Mongolia)" (PDF). American Museum Novitates (3508): 1–16. doi:10.1206/0003-0082(2006)3508[1:EEALSF]2.0.CO;2. S2CID 86032547. Archived (PDF) from the original on 2021-08-31. Retrieved 2020-01-26.
  25. ^ Turner, A. H. (2015). "A Review of Shamosuchus and Paralligator (Crocodyliformes, Neosuchia) from the Cretaceous of Asia". PLOS One. 10 (2) e0118116. Bibcode:2015PLoSO..1018116T. doi:10.1371/journal.pone.0118116. PMC 4340866. PMID 25714338.
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