An assemblage of giant aquatic snakes (Serpentes, Palaeophiidae) from the Eocene of Togo

Swiss Journal of Palaeontology - Tập 140 - Trang 1-18 - 2021
Georgios L. Georgalis1, Guillaume Guinot2, Koffi Evenyon Kassegne3, Yawovi Zikpi Amoudji3, Ampah Kodjo C. Johnson3, Henri Cappetta2, Lionel Hautier2
1Palaeontological Institute and Museum, University of Zurich, Zurich, Switzerland
2Institut Des Sciences de L’Evolution de Montpellier (ISE-M, UMR 5554, CNRS/UM/IRD/EPHE), Université de Montpellier, Montpellier Cedex 5, France
3Département de Géologie, Faculté Des Sciences, Université de Lomé, Lomé, Togo

Tóm tắt

We here describe a monospecific assemblage of giant aquatic snakes from the middle Eocene of Kpogamé, Togo. The material, consisting of large isolated vertebrae, is referred to Palaeophis africanus, an enigmatic palaeophiid species, which was so far otherwise known only from a limited number of vertebrae from the middle Eocene of Nigeria and Angola. Material from the late Eocene of the eastern USA that had been referred to the same species, is here instead considered too fragmentary for species-level determination and Palaeophis africanus is thus so far restricted to Africa. With the aid of micro-CT scanning, we present 3D models of 17 vertebrae, pertaining to different portions of the vertebral column. We provide detailed comparisons of the new material with all named African species of the genus Palaeophis. A tentative diagnosis of Palaeophis africanus is provided. With more than 50 vertebrae, the new Togolese specimens represent the most abundant known material attributed to Palaeophis africanus and significantly enhance our knowledge of the vertebral anatomy and intracolumnar variation for this taxon. Furthermore, this adds to the, as yet, extremely scarce fossil record of squamates from central western Africa, a region where Paleogene herpetofaunas are only rather poorly known.

Tài liệu tham khảo

Andrews, C. W. (1901). Preliminary notes on some recently discovered extinct vertebrates from Egypt. (Part II). Geological Magazine, 8, 436–444. Andrews, C. W. (1906). A descriptive catalogue of the Tertiary Vertebrata of the Fayûm, Egypt, based on the collection of the Egyptian Government in the Geological Museum, Cairo, and on the collection in the British Museum (Natural History), London (p. 324). London: Trustees of the British Museum (Natural History). Andrews, C. W. (1924). Note on some Ophidian vertebræ from Nigeria. Geological Survey of Nigeria, 7, 39–43. Antunes, M. T. (1964). O neocretacico e o cenozoico do litoral de Angola (pp. 1–255). Lisboa: Junta de Investigaçoes do Ultramar. Arambourg, C. (1952). Les vertébrés fossiles des gisements de phosphates (Maroc-Algérie-Tunisie). Notes et Mémoires du Service Géologique du Maroc, 92, 1–372. Berggren, W. A., & Pearson, P. N. (2005). A revised tropical to subtropical Paleogene planktonic foraminiferal zonation. Journal of Foraminiferal Research, 35, 279–298. Bourdon, E., & Cappetta, H. (2012). Pseudo-toothed birds (Aves, Odontopterygiformes) from the Eocene phosphate deposits of Togo, Africa. Journal of Vertebrate Paleontology, 32, 965–970. Cappetta, H., & Traverse, M. (1988). Une riche faune de sélaciens dans le bassin à phosphate de Kpogamé-Hahotoé (Eocène moyen du Togo): Note préliminaire et précisions sur la structure et l’âge du gisement. Geobios, 21, 359–365. Colonna-Cimera, J. (1961). Note sur le démarrage de l’exploitation du gisement de phosphate du bas-Togo. Rapport de la Direction Générale des Mines et de la Géologie, 1961, 1–12. Cope, E. D. (1873). Synopsis of new Vertebrata from the Tertiary of Colorado: obtained during the summer of 1873 (p. 19). Washington D.C.: U.S. Government Printing Office. Folie, A., Mees, F., De Putter, T., & Smith, T. (2021). Presence of the large aquatic snake Palaeophis africanus in the middle Eocene marine margin of the Congo Basin, Cabinda, Angola. Geobios, 66–67, 45–54. Georgalis, G. L., Del Favero, L., & Delfino, M. (2020). Italy’s largest snake: Redescription of Palaeophis oweni from the Eocene of Monte Duello, near Verona. Acta Palaeontologica Polonica, 65, 523–533. Georgalis, G. L., Guinot, G., Kassegne, K. E., Amoudji, Y. Z., Johnson, A. K. C., Cappetta, H., & Hautier, L. (2021). 3D data related to the publication: An assemblage of giant aquatic snakes (Serpentes, Palaeophiidae) from the Eocene of Togo. MorphoMuseuM. https://doi.org/10.18563/journal.m3.154 Georgalis, G. L., & Scheyer, T. M. (2019). A new species of Palaeopython (Serpentes) and other extinct squamates from the Eocene of Dielsdorf (Zurich, Switzerland). Swiss Journal of Geosciences, 112, 383–417. Georgalis, G. L., & Smith, K. T. (2020). Constrictores Oppel, 1811—the available name for the taxonomic group uniting boas and pythons. Vertebrate Zoology, 70, 291–304. Gingerich, P. D., & Cappetta, H. (2014). A new archaeocete and other marine mammals (Cetacea and Sirenia) from lower middle Eocene phosphate deposits of Togo. Journal of Paleontology, 88, 109–129. Head, J. J., Holroyd, P. A., Hutchison, J. H., & Ciochon, R. L. (2005). First report of snakes (Serpentes) from the Late Middle Eocene Pondaung Formation, Myanmar. Journal of Vertebrate Paleontology, 25, 246–250. Hoffstetter, R. (1939). Contribution à l’étude des Elapidæ actuels et fossiles et de l’ostéologie des Ophidiens. Archives du Muséum d’histoire Naturelle de Lyon, 15, 1–78. Hoffstetter, R. (1958). Un serpent marin du genre Pterosphenus (Pt. sheppardi nov. sp.) dans l’Éocène supérieur de l’Equateur (Amérique du Sud). Bulletin de la Société Geologique de France, 8, 45–50. Hoffstetter, R. (1960). Présence de Pterosphenus (serpent paléophidé) dans l’Eocène supèrieur du bord occidental du désert libyque. Comptes Rendus Sommaires de la Société Géologique de France, Paris, 1960, 41. Hoffstetter, R. (1961). Nouvelles récoltes de serpents fossiles dans l’Éocène supérieur de désert Libyque. Bulletin du Muséum National d’histoire Naturelle Paris (série 2), 33, 326–331. Houssaye, A., Herrel, A., Boistel, R., & Rage, J.-C. (2019). Adaptation of the vertebral inner structure to an aquatic life in snakes: Pachyophiid peculiarities in comparison to extant and extinct forms. Comptes Rendus Palevol, 18, 783–799. Houssaye, A., Rage, J.-C., Bardet, N., Vincent, P., Amaghzaz, M., & Meslouh, S. (2013). New highlights about the enigmatic marine snake Palaeophis maghrebianus (Palaeophiidae; Palaeophiinae) from the Ypresian (Lower Eocene) phosphates of Morocco. Palaeontology, 56, 647–661. Hutchison, J. H. (1985). Pterosphenus cf. P. schucherti Lucas (Squamata, Palaeophidae) from the Late Eocene of Peninsular Florida. Journal of Vertebrate Paleontology, 5, 20–23. Janensch, W. (1906). Pterosphenus schweinfurthi Andrews und die Entwicklung der Palaeophiden. Archiv für Biontologie, 1, 307–350. Johnson, A. K. C. (1987). Le Bassin côtier à phosphate du Togo (Maastrichtien-Eocène moyen) (p. 365). Universités de Bourgogne (France) et du Bénin (Togo). Johnson, A. K. C., Rat, P., & Lang, J. (2000). Le bassin sédimentaire à phosphate du Togo (Maastrichtien-Eocene): Stratigraphie, environnements et évolution. Journal of African Earth Sciences, 30, 183–200. Kassegne, K. E., Mourlam, M. J., Guinot, G., Amoudji, Y. Z., Martin, J. E., Togbe, K. A., Johnson, A. K., & Hautier, L. (2021). First partial cranium of Togocetus from Kpogamé (Togo) and the protocetid diversity in the Togolese phosphate basin. Annales de Paléontologie, 107, 102488. Lebrun, R. (2018). MorphoDig, an open-source 3d freeware dedicated to biology. In: IPC5 The 5th International Palaeontological Congress, Paris, France. Linnaeus, C. (1758). Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (p. 824). Stockholm: Laurentii Salvii. Lucas, F. A. (1898). A new snake from the Eocene of Alabama. Proceedings of the United States National Museum, 21, 637–638. Lydekker, R. (1888). Notes on Tertiary Lacertilia and Ophidia. Geological Magazine, 5, 110–113. Massalongo, A. (1859). Specimen photographicum animalium quorumdam plantarumque fossilium agri Veronensis (p. 101). Veronae (= Verona): Vicentinius et Fran chinius Excudebant. McCartney, J. A., & Seiffert, E. R. (2016). A late Eocene snake fauna from the Fayum Depression, Egypt. Journal of Vertebrate Paleontology, 36, e1029580. McCartney, J. A., Roberts, E. M., Tapanila, L., & O’Leary, M. A. (2018). Large palaeophiid and nigerophiid snakes from Paleogene Trans-Saharan Seaway deposits of Mali. Acta Palaeontologica Polonica, 63, 207–220. Nopcsa, F. (1923a). Die familien der reptilien. Fortschritte der Geologie und Paläontologie, 2, 1–210. Nopcsa, F. (1923b). Eidolosaurus und Pachyophis Zwei Neue Neocom-Reptilien. Palaeontographica, 65, 99–154. O’Leary, M., Bouaré, M. L., Claeson, K. M., Hill, R. V., McCartney, J., Sessa, J. A., Sissoko, F., Tapanila, L., Wheeler, E., & Roberts, E. M. (2019). Stratigraphy and paleobiology of the Upper Cretaceous-Lower Paleogene sediments from the trans-Saharan seaway in Mali. Bulletin of the American Museum of Natural History, 436, 1–177. Oppel, M. (1811). Die Ordnungen, Familien und Gattungen der Reptilien als Prodrom einer Naturgeschichte derselben (p. 87). Munich: Joseph Lindauer. Owen, R. (1841). Description of some ophidiolites (Palæophis toliapicus) from the London Clay of Sheppey, indicating an extinct species of serpent. Transaction of the Geological Society Second Series, 6, 209–210. Owen, R. (1850). Part III. Ophidia (Palæophis & c.). In R. Owen (Ed.), Monograph on the fossil Reptilia of the London Clay and of the Bracklesham and other Tertiary beds (pp. 51–63). London: Palæontographical Society of London. Parmley, D., & Case, G. R. (1988). Palaeopheid snakes from the gulf coastal region of North America. Journal of Vertebrate Paleontology, 8, 334–339. Parmley, D., & DeVore, M. (2005). Palaeopheid snakes from the Late Eocene Hardie Mine local fauna of central Georgia. Southeastern Naturalist, 4, 703–722. Parmley, D., & Reed, H. W. (2003). Size and age class estimates of North American Eocene palaeopheid snakes. Georgia Journal of Science, 61, 220–232. Rage, J. C. (1983a). Les serpents aquatiques de l’Éocène européen. Définition des espèces et aspects stratigraphiques. Bulletin du Muséum National d’histoire Naturelle, Section C, Sciences de la Terre, Paléontologie, Géologie, Minéralogie, 4(5), 213–241. Rage, J. C. (1983b). Palaeophis colossaeus nov. sp. (le plus grand Serpent connu?) de l’Eocène du Mali et le problème du genre chez les Palaeopheinae. Comptes Rendus de l’académie des Sciences, Série, 2(296), 1741–1744. Rage, J.-C., Bajpai, S., Thewissen, J. G. M., & Tiwari, B. N. (2003). Early Eocene snakes from Kutch, Western India, with a review of the Palaeophiidae. Geodiversitas, 25, 695–716. Rage, J.-C., & Werner, C. (1999). Mid-Cretaceous (Cenomanian) snakes from Wadi Abu Hashim, Sudan: The earliest snake assemblage. Palaeontologia Africana, 35, 85–110. Rage, J.-C., & Wouters, G. (1979). Decouverte du plus ancien Palaeopheide (Reptilia, Serpentes) dans le Maestrichtien du Maroc. Geobios, 12, 293–296. de Rochebrune, A. T. (1880). Revision des ophidiens fossiles du Museum d’Histoire Naturelle. Nouvelles Archives du Muséum d’Histoire Naturelle 2ème Série, 3, 271–296. Slansky, M. (1962). Contribution à l’étude géologique du bassin sédimentaire côtier du Dahomey et du Togo (p. 335). Thèse université de Nancy. Smith, K. T., & Georgalis, G. L. (In press). The diversity and distribution of Palaeogene snakes: a review, with comments on vertebral sufficiency. In: D. Gower, & H. Zaher (Eds.), The origin and early evolution of snakes. Cambridge: Cambridge University Press. Stromer, E. (1910). Reptilien- und Fischreste aus dem marinen Alttertiär Südtogo (Westafrika). Zeitschrift der Deutschen Geologischen Geselschaft, 62, 478–507. Vandenberghe, N., Speijer, R., & Hilgen, F. J. (2012). The Paleogene period (pp. 855–921). In: F. M. Gradstein, J. G. Ogg, M. Schmitz, & G. M. Ogg (Eds.), The geological time scale 2012. Elsevier Science. Visse, L. (1957). Le faciès phosphate togolais. Rapport inédit (p. 119). Direction Générale des Mines et de la Géologie. Wallach, V., Williams, K. L., & Boundy, J. (2014). Snakes of the world: A catalogue of living and extinct species (p. 1237). Boca Raton: CRC Press. de Zigno, A. (1881). Nuove aggiunte alla fauna eocena del Veneto. Memorie del Reale Istituto Veneto di Scienze, Lettere ed Arti, 21, 775–790. Zouhri, S., Gingerich, P., Khalloufi, B., Bourdon, E., Adnet, S., Jouve, S., Elboudali, N., Amane, A., Rage, J.-C., Tabuce, R., & de Broin, F.L. (2021). Middle Eocene vertebrate fauna from the Aridal Formation, Sabkha of Gueran, southwestern Morocco. In: J.-S. Steyer, M. L. Augé, & G. Métais (Eds.), Memorial Jean-Claude Rage: A life of paleo-herpetologist. Geodiversitas, 43, 121–150. Zouhri, S., Khalloufi, B., Bourdon, E., de Broin, F. L., Rage, J.-C., M’Haïdrat, L., Gingerich, P. D., & Elboudali, N. (2018). Marine vertebrate fauna from the late Eocene Samlat Formation of Ad-Dakhla, south western Morocco. Geological Magazine, 155, 1596–1620. Zvonok, E. A., & Snetkov, P. B. (2012). New findings of snakes of the genus Palaeophis Owen, 1841 (Acrochordoidea: Palaeophiidae) from the middle Eocene of Crimea. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 316, 392–400.