Giải phẫu hộp sọ số hóa của rùa Bắc Mỹ thời Oligocen Stylemys nebrascensis với các bình luận thuế tộc về loài và so sánh với các loài Testudinidae hiện tại thuộc clade Gopherus–Manouria

Serjoscha W. Evers1, Zahra Al Iawati2
1Department of Geosciences, University of Fribourg, Fribourg, Switzerland
2GeoZentrum Nordbayern, Friedrich-Alexander-Universität Erlangen-Nürnberg, Erlangen, Germany

Tóm tắt

Giải phẫu của các loài rùa Bắc Mỹ ít được hiểu biết, mặc dù có một hồ sơ hóa thạch phong phú từ thời Eocene và các tầng trẻ hơn. Stylemys nebrascensis là một loài rùa đáng chú ý trong vấn đề này, vì hàng trăm mẫu vật đã được biết đến từ các trầm tích Oligocen, và loài này là một trong những loài rùa hóa thạch đầu tiên được mô tả trong tài liệu khoa học. Kể từ khi mô tả ban đầu dựa trên một cái mai, nhiều mẫu vật với tư liệu hoàn chỉnh hơn đã được tham chiếu vào Stylemys nebrascensis. Ở đây, chúng tôi xem xét và xác nhận việc tham chiếu một mẫu vật lịch sử quan trọng đến Stylemys nebrascensis, bao gồm vật liệu mai, vật liệu xương không thuộc mai và xương sọ. Điều này cho phép chúng tôi tài liệu hóa các đặc điểm sọ độc đáo của Stylemys nebrascensis (ví dụ, một 'kênh hậu giáp' bất thường kết nối bề mặt sọ phía sau với cavum acustico-jugulare) và tham chiếu một sọ bảo tồn tốt khác đến loài này. Dựa trên quét vi tính hai cái sọ này, chúng tôi cung cấp một mô tả chi tiết về xương sọ và xương hàm của Stylemys nebrascensis. Stylemys nebrascensis có sự kết hợp giữa các đặc điểm sọ cổ nguyên thủy (ví dụ, sự giữ lại của một quá trình jugal giữa) và các đặc điểm tiến hóa được chia sẻ với các loài rùa gopher hiện tại (ví dụ, gờ tiền hàm giữa) cho thấy rằng nó có thể là một đại diện gốc của dòng dõi rùa gopher. Điều này hỗ trợ giả thuyết rằng các loài rùa hiện tại và hóa thạch từ Bắc Mỹ hình thành một nhánh địa lý giới hạn đã phân tách khỏi các họ hàng Châu Á trong kỷ Paleogene.

Từ khóa

#rùa #Stylemys nebrascensis #hóa thạch #giải phẫu #động vật học xương #rùa gopher

Tài liệu tham khảo

Angielczyk, K. D., & Feldman, C. R. (2013). Are diminutive turtles miniaturized? The ontogeny of plastron shape in emydine turtles. Biological Journal of the Linnean Society, 108(4), 727–755. Auffenberg, W. (1964). A redefinition of the fossil tortoise genus Stylemys Leidy. Journal of Paleontology, 38(2), 316–324. Auffenberg, W. (1976). The genus Gopherus (Testudinidae): Pt.1. Osteology and relationships of extant species. Bulletin of the Florida State Museum, Biological Sciences, 20, 47–110. Batsch, A. J.G. C. (1788). Versuch einer Anleitung zur Kenntniss und Geschichte der Thiere und Mineralien. Erster Theil. Allgemeine Geschichte der Natur; besondre der Säugthiere, Vögel, Amphibien und Fische. Jena: Akademischen Buchhandlung. 528 pp. Bels, V., Baussart, S., Davenport, J., Shorten, M., O’Riordan, R. M., Renous, S., & Davenport, J. L. (2007). 8 Functional Evolution of Feeding Behavior in Turtles. Biology of Turtles: From Structures to Strategies of Life. Bramble, D.M. (1971). Functional morphology, evolution and paleoecology of Gopher tortoises [dissertation]. Berkeley: University of California. 341 pp. Bramble, D. M. (1974). Occurrence and significance of the os transiliens in gopher tortoises. Copeia. https://doi.org/10.2307/1443010 Bramble, D.M., & Wake, D.B. (1985). Feeding mechanisms of lower tetrapods, in Functional Vertebrate Morphology, M. Hildebrand, D.M. Bramble, K.F. Liem, and D.B. Wake (eds.), Cambridge: Harvard University Press, 230. https://doi.org/10.4159/harvard.9780674184404.c13 Brinkman, D., Hart, M., Jamniczky, H., & Colbert, M. (2006). Nichollsemys baieri gen. et sp. nov, a primitive chelonioid turtle from the late Campanian of North America. Paludicola, 5(4), 111–124. Case, E. C. (1920). Note on a specimen of Stylemys nebraskensis Leidy. American Journal of Science, 47, 435–437. Case, E. C. (1925). A specimen of Stylemys nebrascensis Leidy, with the skull preserved. Contributions from the Museum of Paleontology of the University of Michigan, 2(4), 87–91. Case, E. C. (1936). A specimen of Stylemys nebrascensis Leidy, showing the bones of the feet and limbs. Contributions from the Museum of Paleontology of the University of Michigan, 5(6), 69–73. Cope, E. D. (1884): The Vertebrata of the Tertiary formations of the West. In: Report of the United States Geological Survey of the Territories, Volume 3. Government Printing Office; pp. 1–1009. Crumly, C. R. (1984). A hypothesis for the relationships of land tortoise genera (Family Testudinidae). Studia Palaeocheloniologica, 1, 115–124. Crumly, C. R. (1994). Phylogenetic systematics of North American tortoises (genus Gopherus): Evidence for their classification. Fish and Wildlife Research, 13, 7–32. CTEES. (2022a). University of Michigan Museum of Paleontology, CTEES. (2022). CT Data of UMMP VP 9318, Stylemys nebrascensis skull [Data set], University of Michigan—Deep Blue Data. https://doi.org/10.7302/74pd-kb09 CTEES. (2022b). University of Michigan Museum of Paleontology, CTEES. (2022). CT Data of UMMP VP 9318, Stylemys nebrascensis postcrania (cervical vertebrae, L+R humerus, R scapula, R coracoid, L femur) [Data set], University of Michigan - Deep Blue Data. doi: https://doi.org/10.7302/h3aw-9427 De Stefano, G. (1902). Stylemys Bottii n. f. Rivista Italiana Di Paleontologia, 8, 72–76. Ernst, C. H., & Barbour, R. W. (1989). Turtles of the World (p. 313). Smithsonian Institution Press. Evers, S.W. (2021). Fossil turtle cranial CT scans. MorphoSource. Retrieved from https://www.morphosource.org/projects/000350496 Evers, S. W., & Al Iawati, Z. (2023). 3D models of Stylemys nebrascensis and Manouria impressa. MorphoSource. Retrieved from https://www.morphosource.org/projects/000516064 Evers, S. W., Barrett, P. M., & Benson, R. B. (2019a). Anatomy of Rhinochelys pulchriceps (Protostegidae) and marine adaptation during the early evolution of chelonioids. PeerJ, 7, e6811. https://doi.org/10.7717/peerj.6811 Evers, S. W., & Benson, R. B. (2018). Project : Evers & Benson 2018, turtle CT data and 3D models. MorphoSource. Retrieved from http://www.morphosource.org/Detail/ProjectDetail/Show/project_id/462 Evers, S. W., & Benson, R. B. (2019). A new phylogenetic hypothesis of turtles with implications for the timing and number of evolutionary transitions to marine lifestyles in the group. Palaeontology, 62(1), 93–134. https://doi.org/10.1111/pala.12384 Evers, S. W., & Joyce, W. G. (2020). A re-description of Sandownia harrisi (Testudinata: Sandownidae) from the Aptian of the Isle of Wight based on based on computed tomography scans. Royal Society Open Science, 7(2), 191936. https://doi.org/10.1098/rsos.191936 Evers, S. W., Joyce, W. G., Choiniere, J. N., Ferreira, G. S., Foth, C., Hermanson, G., Hongyu, Y., Johnson, C. M., Werneburg, I., & Benson, R. B. (2022). Independent origin of large labyrinth size in turtles. Nature Communications, 13(1), 5807. https://doi.org/10.1038/s41467-022-33091-5 Evers, S. W., Neenan, J. M., Ferreira, G. S., Werneburg, I., Barrett, P. M., & Benson, R. B. (2019b). Neurovascular anatomy of the protostegid turtle Rhinochelys pulchriceps and comparisons of membranous and endosseous labyrinth shape in an extant turtle. Zoological Journal of the Linnean Society, 187(3), 800–828. https://doi.org/10.1093/zoolinnean/zlz063 Evers, S. W., & Ponstein, J. (2022). Turtle mandibular anatomy and 3D models. MorphoSource. Retrieved from https://www.morphosource.org/projects/000408332 Evers, S. W., Ponstein, J., Jansen, M. A., Gray, J. A., & Froebisch, J. (2023). A systematic compendium of turtle mandibular anatomy using digital dissections of soft tissue and osteology. The Anatomical Record, 306(6), 1228–1303. https://doi.org/10.1002/ar.25037 Evers, S. W., Rollot, Y., & Joyce, W. G. (2020). Cranial osteology of the Early Cretaceous turtle Pleurosternon bullockii (Paracryptodira: Pleurosternidae). PeerJ, 8, e9454. https://doi.org/10.7717/peerj.9454 Evers, S. W., Rollot, Y., & Joyce, W. G. (2021). New interpretation of the cranial osteology of the Early Cretaceous turtle Arundelemys dardeni (Paracryptodira) based on a CT-based re-evaluation of the holotype. PeerJ, 9, e11495. https://doi.org/10.7717/peerj.11495 Ferreira, G. S., Werneburg, I., Lautenschlager, S., & Evers, S. W. (2022). Contrasting brains and bones: neuroanatomical evolution of turtles (Testudinata). in Paleoneurology of amniotes: new directions in the study of fossil endocasts (pp. 79–121). Springer International Publishing. https://doi.org/10.1007/978-3-031-13983-3_4 Fish, J. F., & Stayton, C. T. (2014). Morphological and mechanical changes in juvenile red-eared slider turtle (Trachemys scripta elegans) shells during ontogeny. Journal of Morphology, 275(4), 391–397. Fitzinger, L. J. (1826). Neue Classification der Reptilien nach ihren natürlichen Verwandtschaften (p. 66). Verlag J. G. Heubner. Gaffney, E. S. (1972). An illustrated glossary of turtle skull nomenclature. American Museum Novitates, 2486, 1–33. Gaffney, E. S. (1979). Comparative cranial morphology of Recent and fossil turtles. Bulletin of the American Museum of Natural History, 164(2), 67–376. Gaffney, E. S., Tong, H., & Meylan, P. A. (2006). Evolution of the side-necked turtles: the families Bothremydidae, Euraxemydidae, and Araripemydidae. Bulletin of the American Museum of Natural History, 300, 1–698. Georgalis, G. L., Macaluso, L., & Delfino, M. (2021). A review of the fossil record of Afro-Arabian turtles of the clade Testudinoidea. Bulletin of the Peabody Museum of Natural History, 62, 43–78. https://doi.org/10.3374/014.062.0103 Georgi, J. A., & Sipla, J. S. (2008). Comparative and functional anatomy of balance in aquatic reptiles and birds. Sensory Evolution on the Threshold: Adaptations in Secondarily Aquatic Vertebrates, 15, 233–256. https://doi.org/10.1525/california/9780520252783.003.0015 Gilmore, C. W. (1946). The osteology of the fossil turtle Testudo praeextans Lambe, with notes on other species of Testudo from the Oligocene of Wyoming. Proceedings of the United States National Museum, 96, 293–310. Gray, J. E. (1825). A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Annals of Philosophy, 10, 193–217. Hay, O. P. (1908). The Fossil Turtles of North America. Washington, DC: Carnegie Institution of Washington. (Publications 75) 568 pp. Hutchison, J. H. (1996). Testudines. In D. R. Prothero & R. J. Emry (Eds.), The terrestrial Eocene-Oligocene transition in North America (pp. 337–353). Cambridge University Press. Hutchison, J. H., & Bramble, D. M. (1981). Homology of the plastral scales of the Kinosternidae and related turtles. Herpetologica, 73(2), 73–85. Jones, M. E., Werneburg, I., Curtis, N., Penrose, R., O’Higgins, P., Fagan, M. J., & Evans, S. E. (2012). The head and neck anatomy of sea turtles (Cryptodira: Chelonioidea) and skull shape in Testudines. PLoS ONE, 7(11), e47852. https://doi.org/10.1371/journal.pone.0047852 Joyce, W. G., Anquetin, J., Cadena, E.-A., Claude, J., Danilov, I. G., Evers, S. W., Fer- reira, G. S., Gentry, A. D., Georgalis, G. L., Lyson, T. R., Pérez-García, A., Rabi, M., Sterli, J., Vitek, N. S., & Parham, J. F. (2021). A nomenclature for fossil and living turtles using phylogenetically defined clade names. Swiss Journal of Paleontology, 140(5), 1–45. https://doi.org/10.1186/s13358-020-00211-x Joyce, W. G., & Bell, C. J. (2004). A review of the comparative morphology of extant testudinoid turtles (Reptilia: Testudines). Asiatic Herpetological Research, 10, 53. Lambe, L. M. (1913). Description of a new species of Testudo, and of a remarkable specimen of Stylemys Nebrascensis, from the Oligocene of Wyoming. The Ottawa Naturalist, 14, 57–63. Leidy, J. (1851). Description of a new fossil tortoise from Nebraska territory. Proceedings of the Academy of Natural Sciences of Philadelphia, 5, 172–173. Leidy, J. (1853). A description of remains of extinct mammalia and chelonian from the Mauvaises Terres of Nebraska (p. 119). Smithsonian Contributions to Knowledge. Lemell, P., Natchev, N., Beisser, C. J., & Heiss, E. (2019). Feeding in turtles: understanding terrestrial and aquatic feeding in a diverse but monophyletic group. Feeding in Vertebrates: Evolution, Morphology, Behavior, Biomechanics. https://doi.org/10.1007/978-3-030-13739-7_16 Meylan, P. A., & Sterrer, W. (2000). Hesperotestudo (Testudines: Testudinidae) from the Pleistocene of Bermuda, with comments on the phylogenetic position of the genus. Zoological Journal of the Linnean Society, 128(1), 51–76. https://doi.org/10.1111/j.1096-3642.2000.tb00649.x Natchev, N., Tzankov, N., Werneburg, I., & Heiss, E. (2015). Feeding behaviour in a ‘basal’ tortoise provides insights on the transitional feeding mode at the dawn of modern land turtle evolution. PeerJ, 3, e1172. https://doi.org/10.7717/peerj.1172 Neenan, J. M., Reich, T., Evers, S. W., Druckenmiller, P. S., Voeten, D. F., Choiniere, J. N., Barret, P. M., Pierce, S. E., & Benson, R. B. (2017). Evolution of the sauropterygian labyrinth with increasingly pelagic lifestyles. Current Biology, 27(24), 3852–3858. https://doi.org/10.1016/j.cub.2017.10.069 Pritchard, P. C. H. (1988). A survey of neural bone variation among recent chelonian species, with functional interpretations. Acta Zoologica Cracoviensia, 31(2), 11–27. Rabbitt, R. D., Damiano, E. R., & Grant, J. W. (2004). Biomechanics of the semicircular canals and otolith organs. In:The vestibular system (pp. 153–201). Springer New York. https://doi.org/10.1007/0-387-21567-0_4 Rabi, M., Zhou, C. F., Wings, O., Ge, S., & Joyce, W. G. (2013). A new xinjiangchelyid turtle from the Middle Jurassic of Xinjiang, China and the evolution of the basipterygoid process in Mesozoic turtles. BMC Evolutionary Biology, 13(1), 1–29. https://doi.org/10.1186/1471-2148-13-203 Raselli, I. (2018). Comparative cranial morphology of the Late Cretaceous protostegid sea turtle Desmatochelys lowii. PeerJ, 6, e5964. https://doi.org/10.7717/peerj.5964 Ray, C. E. (1959). A sesamoid bone in the jaw musculature of Gopherus polyphemus (Reptilia: Testudiniae). Anatomischer Anzeiger, 107, 85–91. Rollot, Y., Evers, S. W., & Joyce, W. G. (2021a). A review of the carotid artery and facial nerve canal systems in extant turtles. PeerJ, 8, e10475. https://doi.org/10.7717/peerj.10475 Rollot, Y., Evers, S. W., & Joyce, W. G. (2021b). A redescription of the Late Jurassic (Tithonian) turtle Uluops uluops and a new phylogenetic hypothesis of Paracryptodira. Swiss Journal of Palaeontology, 140(1), 1–30. https://doi.org/10.1186/s13358-021-00234-y Schwab, J. A., Young, M. T., Neenan, J. M., Walsh, S. A., Witmer, L. M., Herrera, Y., Allain, R., Brochu, C. A., Choiniere, J. N., Clark, J. M., Dollman, K. N., Etches, S., Fritsch, G., Gignac, P. M., Ruebenstahl, A., Turner, A. H., Vignaud, P., Wilberg, E. W., Xu, X., … Brusatte, S. L. (2020). Inner ear sensory system changes as extinct crocodylomorphs transitioned from land to water. Proceedings of the National Academy of Sciences, 117(19), 10422–10428. https://doi.org/10.1073/pnas.2002146117 Shiino, K. (1912). Beitrag zur Kenntnis der Gehirnnerven der Schildkröten. Anatomische Hefte, 47, 1–34. Smith, L. L., Pedrono, M., Dorazio, R. M., & Bishko, J. (2001). Morphometrics, sexual dimorphism, and growth in the Angonoka tortoise (Geochelone yniphora) of western Madagascar. African Journal of Herpetology, 50(1), 9–18. Sterli, J., Müller, J., Anquetin, J., & Hilger, A. (2010). The parabasisphenoid complex in Mesozoic turtles and the evolution of the testudinate basicranium. Canadian Journal of Earth Sciences, 47(10), 1337–1346. https://doi.org/10.1139/E10-061 Strömberg, C. A. (2004). Using phytolith assemblages to reconstruct the origin and spread of grass-dominated habitats in the great plains of North America during the late Eocene to early Miocene. Palaeogeography, Palaeoclimatology, Palaeoecology, 207(3–4), 239–275. Szalai, T. (1930). Bionomische und methodologisch-systematische Untersuchungen an rezenten und fossilen Testudinaten. Palaeobiologica, 3, 347–364. Thomson, R. C., Spinks, P. Q., & Shaffer, H. B. (2021). A global phylogeny of turtles reveals a burst of climate-associated diversification on continental margins. Proceedings of the National Academy of Sciences. https://doi.org/10.1073/pnas.2012215118 TTWG Turtle Taxonomy Working Group, Rhodin, A. G. J., Iverson, J. B., Bour, R., Fritz, U., Georges, A., Shaffer, H. B., & van Dijk, P. P. (2021). Turtles of the world, 9th edition: Annotated checklist of taxonomy, synonymy, distribution, and conservation status. Chelonian Research Monographs, 8, 1–472. https://doi.org/10.3854/crm.8.checklist.atlas.v9.2021 Vlachos, E. (2018). A review of the fossil record of North American turtles of the clade Pan-Testudinoidea. Bulletin of the Peabody Museum of Natural History, 59(1), 3–94. https://doi.org/10.3374/014.059.0101 Vlachos, E., & Rabi, M. (2018). Total evidence analysis and body size evolution of extant and extinct tortoises (Testudines: Cryptodira: Pan-Testudinidae). Cladistics, 34(6), 652–683. https://doi.org/10.1111/cla.12227 Werneburg, I., & Joyce, W. G. (2021). Cranial Turtle CT scans. MorphoSource. Retrieved from https://www.morphosource.org/projects/000353832 Williams, E. E. (1950). Testudo cubensis and the evolution of western hemisphere tortoises. Bulletin of the American Museum of Natural History, 95, 7–36. Williams, E. E. (1952). A new fossil tortoise from Mona Island, West-Indies, and a tentative arrangement of the tortoises of the world. Bulletin of the American Museum of Natural History, 99(9), 545–560.