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Axial Skeleton of Subadult Protoceratops andrewsi from Djadokhta Formation (Upper Cretaceous, Mongolia)

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Abstract

Protoceratops andrewsi Granger and Gregory, 1923, is one of the most famous Upper Cretaceous horned dinosaurs of Central Asia. It was described from a separate skull found at Bayan Zag locality (Djadokhta formation, Mongolia). The holotype axial skeleton of this species is unknown, and its description appeared to be rather aggregated, capturing structural features of most ceratopsians from this locality. Polymorphism determined in this species is normally explained by gender, age and individual dimorphism. However, there is an opinion that P. andrewsi is a combined taxon that includes representatives of several species, but it is as yet difficult to determine the number of these species using the skull material. The study of the complete skeleton of P. andrewsi ZPAL Mg D-II/3 from the collection of Kozłowski Institute of Paleobiology, Polish Academy of Sciences (Warsaw, Poland), found at Bayan Zag, provided a solution to this problem. A detailed description of ZPAL Mg D-II/3 vertebrae and the study of its axial skeleton, taking into account its age and gender, revealed that this sample, most likely, was a large immature female. Comparative analysis of the vertebrae of this specimen with identical (of the same index number) vertebrae of specimens without associated skulls (PIN 3143/9 and PIN 3143/16) from the age-matched Tögrögiin Shiree locality (Mongolia), which are housed at Borissiak Paleontological Institute, Russian Academy of Sciences (Moscow, Russia), convinced us that they belong to the same morphotype. This makes it possible to diagnose P. andrewsi from vertebrae considering the age and gender of compared specimens.

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REFERENCES

  1. Alifanov, V.R., Two new dinosaurs of the infraorder Neoceratopsia (Ornithischia) from the Upper Cretaceous of the Nemegt Depression, Mongolian People’s Republic, Paleontol. J., 2003, vol. 37, no. 5, pp. 524–534.

    Google Scholar 

  2. Alifanov, V.R., The tiny horned dinosaur Gobiceratops minutus gen. et sp. nov. (Bagaceratopidae, Neoceratopsia) from the Upper Cretaceous of Mongolia, Paleontol. J., 2008, vol. 42, no. 6, pp. 621–633.

    Article  Google Scholar 

  3. Alifanov, V.R., Infraorder Ornithischia, in Iskopaemye reptilii i ptitsy (Fossil Reptiles and Birds), Kurochkin, E.N., and Lopatin, A.V., Eds., Moscow: GEOS, 2012, part 2, pp. 241–309.

  4. Bohlin, B., Fossil reptiles from Mongolia and Kansu, Sino-Swed. Exped. Publ., 1953, vol. 37, pp. 1–113.

    Google Scholar 

  5. Brochu, C.A., Closure of neurocentral sutures during crocodilian ontogeny: implications for maturity assessment in fossil archosaurs, J. Vertebr. Paleontol., 1996, vol. 16, no. 1, pp. 49–62.

    Article  Google Scholar 

  6. Brovar, V.Ya., Biomechanics of the withers (in connection with the issue of the role of spinous processes in vertebrates), Tr. Mosk. Zootekhn. Inst., 1935, vol. 2, pp. 217–251.

    Google Scholar 

  7. Brown, B. and Schlaikjer, E.M., The structure and relationship of Protoceratops, Ann. N.Y. Acad. Sci., 1940, vol. 40, pp. 133–265.

    Article  Google Scholar 

  8. Brown, B. and Schlaikjer, E.M., The skeleton of Leptoceratops with the description of a new species, Am. Mus. Novit., 1942, vol. 1169, pp. 1–15.

    Google Scholar 

  9. Carroll, R.L., Vertebrate Paleontology and Evolution, New York: Freeman, 1988.

    Google Scholar 

  10. Chinnery, B.J. and Weishampel, D.B., Montanoceratops cerorhynchus (Dinosauria: Ceratopsia) and relationships among basal Neoceratopsia, J. Vertebr. Paleontol., 1998, vol. 18, no. 3, pp. 569–585.

    Article  Google Scholar 

  11. Chionidi, R.J. and Nielsen, S.W., Vertebral osteophytes in an iguanid lizard, Vertebr. Pathol., 1983, vol. 20, no. 3, pp. 372–375.

    Google Scholar 

  12. Coombs, W.P., Juvenile ceratopsian from Mongolia—the smallest known dinosaur specimens, Nature, 1980, vol. 283, pp. 380–381.

    Article  Google Scholar 

  13. Coombs, W.P., Juvenile specimens of the ornithischian dinosaur Psittacosaurus, Palaeontology, 1982, vol. 25, pp. 89–107.

    Google Scholar 

  14. Filler, A.G., Axial character seriation in mammals: an historical and morphological exploration of the origin, development, use, and current collapse of the homology paradigm, Ph.D. Thesis, Cambridge, Massachusetts: Harvard University, Department of Anthropology, 2007.

  15. Gilmore, C.W., Brachyceratops, a ceratopsian dinosaur from the two medicine formation of Montana, with notes on associated fossil reptiles, United States Geological Survey Washington, Professional Paper, 1917, vol. 103, pp. 1–45.

    Google Scholar 

  16. Granger, W.K. and Gregory, W.K., Protoceratops andrewsi, a pre-ceratopsian dinosaur from Mongolia, Am. Mus. Novit., 1923, vol. 72, pp. 1–6.

    Google Scholar 

  17. Gregory, W.K. and Mook, C.C., On Protoceratops, a primitive ceratopsian dinosaur from the Lower Cretaceous of Mongolia, Am. Mus. Novit., 1925, vol. 156, pp. 1–9.

  18. Haines, R.W., Epiphyses and sesamoids, in Biology of the Reptilia, Gans, C., d’A. Bellairs, A., and Parsons, T.S., Eds., London, 1969, vol. 1, pp. 81–115.

  19. Handa, N., Watabe, V., and Tsogtbaatar, K., New specimens of Protoceratops (Dinosauria: Neoceratopsia) from Upper Cretaceous in Udyn Sayr, Southern Gobi area, Mongolia, Paleontol. Res., 2012, vol. 16, no. 3, pp. 179–198.

    Article  Google Scholar 

  20. Hoffstetter, R. and Gasc, J.-P., Vertebrae and ribs of modern reptiles, in Biology of the Reptilia, Gans, C., d’A. Bellairs, A., and Parsons, T.S., Eds., London, 1969, vol. 1, pp. 201–335.

  21. Klimov, A.F., Anatomiya domashnikh zhivotnykh (Anatomy of Domestic Animals), Moscow: Gos. S.-Kh. Lit., 1950, vol. 1.

  22. Kovalenko, E.E., Anomalii pozvonochnika beskhvostykh amfibii (Anomalies of the Spine of Anura), St. Petersburg: St.‑Peterb. Univ., 1992.

  23. Kurzanov, S.M., On sexual dimorphism of Protoceratops, Paleontol. Zh., 1972, no. 1, pp. 104–112.

  24. Kuznetsov, A.N. and Tereshchenko, V.S., A method for estimation of lateral and vertical mobility of platycoelous vertebrae of tetrapods, Paleontol. J., 2010, vol. 44, no. 2, pp. 209–225.

    Article  Google Scholar 

  25. Lambert, O., Godefroit, P., Li, H., et al., A new species of Protoceratops (Dinosauria, Neoceratopsia) from the Late Cretaceous of Inner Mongolia (P.R. China), Bull. Inst. R. Sci. Nat. Belg., Sci. Terre, Suppl., 2001, vol. 71, pp. 5–28.

    Google Scholar 

  26. Lange, N.O., The correlation between the reduction of the extremities and the development of the abdominal muscles in reptiles, À la mèmoire de A.N. Sewertzoff 1866–1936 (Recueil des travaux publiè en deux volumes), Moscou: Institut A.N. Sewertzoff de Morphologie Ѐvolutive, Éditions de L’Académie des Sciences de L’URSS, 1940, vol. 2, no. 1, pp. 349–374.

    Google Scholar 

  27. Makovicky, P.J. and Norell, M.A., Yamaceratops dorngobiensis, a new primitive ceratopsian (Dinosauria: Ornithischia) from the Cretaceous of Mongolia, Am. Mus. Novit., 2006, vol. 3530, pp. 1–42.

    Article  Google Scholar 

  28. Maryańska, T. and Osmólska, H., Protoceratopsidae (Dinosauria) of Asia, Palaeontol. Pol., 1975, vol. 33, pp. 133–182.

    Google Scholar 

  29. Niedźwiedzki, G., Singer, T., Gierliński, G.D., and Lockley, M.G., A protoceratopsid skeleton with an associated track from the Upper Cretaceous of Mongolia, Cretaceous Res., 2012, vol. 33, pp. 7–10.

    Article  Google Scholar 

  30. Romer, A.S., Osteology of the Reptiles, Chicago: Univ. Chicago Press, 1956, pp. 79–89.

    Google Scholar 

  31. Sereno, P.C., Xijin, Z., Brown, L., and Lin, T., New psittacosaurid highlights skull enlargement in horned dinosaurs, Acta Palaeontol. Pol., 2007, vol. 52, no. 2, pp. 275–284.

    Google Scholar 

  32. Słowiak, J., Tereshchenko, V.S., and Fostowicz-Frelik, Ł., Appendicular skeleton of Protoceratops andrewsi (Dinosauria, Ornithischia): comparative morphology, ontogenetic changes, and the implications for stem ceratopsian locomotion, PeerJ, 2019, no. 7. https://doi.org/10.7717/peerj.7324

  33. Sternberg, C.M., Complete skeleton of Leptoceratops gracilis Brown from the Upper Edmonton Member on Red Deer River, Alberta, Bull. Natl. Mus. Can., 1951, vol. 123, pp. 225–255.

    Google Scholar 

  34. Sukhanov, V.B. and Tereshchenko, V.S., Composition and formation of the sacrum of dinosaurs, in Mezhdunar. Konf. “Problemy paleontologii Tsentral’noi Azii (k 35-letiyu Sovmestnoi Rossiisko-Mongol’skoi paleontologicheskoi ekspeditsii),” 27–28 maya 2004 g., Tezisy dokladov (Int. Conf. “Problems of Paleontology of Central Asia (to the 35th Anniversary of the Joint Russian–Mongolian Paleontological Expedition),” May 27–28, 2004), Moscow: Paleontol. Inst. Ross. Akad. Nauk, 2004, pp. 56–58.

  35. Tereshchenko, V.S., Vertebra size restoration technique in fossil tetrapods, Paleontol. Zh., 1990, no. 4, pp. 98–102.

  36. Tereshchenko, V.S., Towards the reconstruction of the spinal column of the Protoceratops, Paleontol. Zh., 1991a, no. 2, pp. 86–96.

  37. Tereshchenko, V.S., Sexual dimorphism of the postcranial skeleton of lizards of the family Agamidae (Lacertilia), Zool. Zh., 1991b, vol. 70, no. 10, pp. 91–103.

    Google Scholar 

  38. Tereshchenko, V.S., Sexual dimorphism in the postcranial skeleton of protoceratopsids (Neoceratopsia, Protoceratopsidae) from Mongolia, Paleontol. J., 2001, vol. 35, no. 4, pp. 415–425.

    Google Scholar 

  39. Tereshchenko, V.S., On the heterocelous vertebrae in horned dinosaurs (Protoceratopidae, Neoceratopsia), Paleontol. J., 2004a, vol. 38, no. 2, pp. 200–205.

    Google Scholar 

  40. Tereshchenko, V.S., The morphology of the axial skeleton of the horned dinosaurs (Neoceratopsia: Protoceratopoidea) from the Upper Cretaceous of Mongolia, Extended Abstract of Cand. Sci. (Biol.) Dissertation, Moscow: Paleontol. Inst. Ross. Akad. Nauk, 2004b.

  41. Tereshchenko, V.S., Key to protoceratopoid vertebrae (Ceratopsia, Dinosauria) from Mongolia, Paleontol. J., 2007, vol. 41, no. 2, pp. 175–188.

    Article  Google Scholar 

  42. Tereshchenko, V.S., Adaptive features of Protoceratopoids (Ornithischia: Neoceratopsia), Paleontol. J., 2008, vol. 42, no. 3, pp. 273–286.

    Article  Google Scholar 

  43. Tereshchenko, V.S., Methods for size reconstruction in distorted and damaged vertebrae of fossil tetrapods, Paleontol. J., 2015, vol. 49, no. 3, pp. 293–303.

    Article  Google Scholar 

  44. Tereshchenko, V.S., On polymorphism of Proto-ceratops andrewsi Granger et Gregory, 1923 (Protoceratopidae, Neoceratopsia), Paleontol. J., 2018, vol. 52, no. 4, pp. 429–444.

    Article  Google Scholar 

  45. Tereshchenko V.S., Sexual dimorphism in the postcranial skeleton of dinosaurs, Paleontol. J., 2020, vol. 54, no. 12, pp. 1410–1433.

    Article  Google Scholar 

  46. Tereshchenko, V.S. and Alifanov, V.R., Bainoceratops efremovi, a new protoceratopid dinosaur (Protoceratopidae, Neoceratopsia) from the Bain-Dzak locality (South Mongolia), Paleontol. J., 2003, vol. 37, no. 3, pp. 293–302.

    Google Scholar 

  47. Tereshchenko, V.S. and Singer, T., Structural features of neural spines of the caudal vertebrae of protoceratopoids (Ornithischia: Neoceratopsia), Paleontol. J., 2013, vol. 47, no. 6, pp. 618–630.

    Article  Google Scholar 

  48. Tereshchenko, V.S. and Sukhanov, V.B., Variability of the axial skeleton of protoceratopoids of Central Asia, in Mezhdunar. konf. k 40-letiyu Sovmestnoi rossiisko-mongol’skoi paleontologicheskoi ekspeditsii “Paleontologiya Tsentral’noi Azii,” 18–19 noyabrya 2009, Tezisy dokladov (Int. Conf. to the 40th Anniversary of the Joint Russian–Mongolian Paleontological Expedition “Paleontology of Central Asia”, November 18–19, 2009, Abstracts of Papers), Moscow: Paleontol. Inst. Ross. Akad. Nauk, 2009, pp. 83–86.

  49. Tereshchenko, V.S. and Sukhanov, V.B., Study of the axial skeleton of protoceratopoids of Mongolia: approaches and methods, in Mezhdunar. konf. k 45-letiyu Sovmestnoi rossiisko-mongol’skoi paleontologicheskoi ekspeditsii “Paleontologiya Tsentral’noi Azii i sopredel’nykh regionov,” 12–13 noyabrya 2014, Tezisy dokladov (Int. Conf. to the 45th Anniversary of the Joint Russian–Mongolian Paleontological Expedition “Paleontology of Central Asia and Adjacent Regions,” November 12–13, 2014, Abstracts of Papers), Moscow: Paleontol. Inst. Ross. Akad. Nauk, 2014, pp. 72–75.

  50. Tsuihiji, T. and Makovicky, P.J., Homology of the neoceratopsian cervical bar elements, J. Paleontol., 2007, vol. 81, no. 5, pp. 1132–1138.

    Article  Google Scholar 

  51. Vygodskii, M.Ya., Spravochnik po elementarnoi matematike (Handbook of Elementary Mathematics), Moscow: Nauka, 1975, pp. 1–416.

  52. Xing, L., Rothschild, B.M., Ran, H., et al., Vertebral fusion in two Early Jurassic sauropodomorph dinosaurs from the Lufeng formation of Yunnan, China, Acta Palaeontol. Pol., 2015, vol. 60, no. 3, pp. 643–649.

    Google Scholar 

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ACKNOWLEDGMENTS

The author thanks T. Singer (University of Warsaw, Poland) for the technical processing of this material, without which this research would not have been possible, and M. Borsuk-Bialynicka and J. Dzik (Institute of Paleobiology, PAN, Poland) for their active support. Many thanks to M. Dziewiński (Institute of Paleobiology PAN, Poland) for the photographs he prepared for this work, as well as to V.B. Sukhanov and A.G. Sennikov (PIN RAS) for reviewing the manuscript, valuable advice, and comments.

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This study was supported by the Russian Foundation for Basic Research (project no. 20-04-00070).

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Correspondence to V. S. Tereshchenko.

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Translated by M. Batrukova

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Tereshchenko, V.S. Axial Skeleton of Subadult Protoceratops andrewsi from Djadokhta Formation (Upper Cretaceous, Mongolia). Paleontol. J. 55, 1408–1457 (2021). https://doi.org/10.1134/S0031030121120030

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