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A Mandible of the Hyracoid Mammal Titanohyrax

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A Mandible of the Hyracoid Mammal Titanohyrax A mandible of the hyracoid mammal Titanohyrax andrewsi in the collections of the Muséum National d’Histoire Naturelle, Paris (France) with a reassessment of the species Rodolphe Tabuce To cite this version: Rodolphe Tabuce. A mandible of the hyracoid mammal Titanohyrax andrewsi in the collections of the Muséum National d’Histoire Naturelle, Paris (France) with a reassessment of the species. Palaeover- tebrata, Association Palaeovertebrata., 2016, 40 (1), pp.e4. 10.18563/pv.40.1.e4. hal-03100188 HAL Id: hal-03100188 https://hal.archives-ouvertes.fr/hal-03100188 Submitted on 21 Jan 2021 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. ARTICLE A mandible of the hyracoid mammal Titanohyrax andrewsi in the collections of the Muséum National d’Histoire Naturelle, Paris (France) with a reassessment of the species RODOLPHE TABUCE Institut des Sciences de l’Évolution (UM, CNRS, IRD, EPHE), c.c. 064, Université de Montpellier, Place Eugène Bataillon, 34095 Montpellier cedex 05, France Email: [email protected] Abstract: An unpublished mandible of the large hyracoid Titanohyrax andrewsi from the early Oligocene Jebel Qatrani Formation, Fayum Depression, Egypt is described. This specimen has a twofold importance. Firstly, it opens an unexpected window on early paleontological research in the Fayum because it was discovered as early as 1904 by the French paleontologist René Fourtau during an expedition to the Fayum organized by the Muséum National d’Histoire Naturelle, Paris (MNHN). This expedition has remarkably never been mentioned in the literature. Secondly, the mandible documents the best-preserved specimen of T. andrewsi, permitting a revision of one of the very rare Paleogene hyracoids. Interestingly, the new mandible was discovered two years before the first report of the species by Charles W. Andrews. The hypodigm of T. andrewsi is reviewed and the dentition as a whole is compared in detail, notably with other Titanohyrax species from the Fayum. The validity of the large Titanohyrax “schlosseri” species is discussed, but a pronounced sexual size dimorphism for T. andrewsi is favoured. Keywords: Titanohyracidae, Oligocene, Afro-Arabia, Fayum Submitted 1 December 2015, Accepted 15 March 2016 Published Online 18 April 2016, doi: 10.18563/pv.40.1.e4 © Copyright Rodolphe Tabuce April 2016 INTRODUCTION yet undescribed, ?Titanohyrax species are known from the Glib Zegdou Formation (Tabuce et al., 2011) (Fig. 1). Besides its size and relatively high specific diversity, the genus Titanohyrax is The hyracoid Titanohyrax is one of the most iconic mammals peculiar among the numerous Paleogene hyracoids by its lo- from the African Paleogene fauna since Matsumoto (1922:844) phoselenodont dental pattern, fully molariform premolars, and described T. ultimus from the early Oligocene of the Jebel relatively high-crowned cheek teeth (with the exception of T. Qatrani Formation, Fayum Depression, Egypt, as an “extremely mongereaui). This morphology, which is also characteristic of gigantic species, being the largest of all the hyracoids hitherto Antilohyrax ― another titanohyracid from the Fayum ―, is known”. Body mass estimates for this species range from indicative of a probable folivorous diet (Rasmussen, 1989). around 600 kg to more than 1300 kg, close in weight to the extant Sumatran rhinoceros (Schwartz et al., 1995, but see Despite their apparent central role in the Paleogene Pickford 2015:204). The large size of T. ultimus evidently mammalian communities, the various Titanohyrax species are contrasts with the small sizes of the three living hyracoid genera still poorly known due to their rarity in the fossil record. It is whose body masses range from 1.3 to 5.5 kg (Shoshani et al., worth mentioning here that T. mongereaui is only documented 2013). Besides T. ultimus, three other Titanohyrax species by its holotype despite recent intensive field research in the are known from the Jebel Qatrani Formation: T. angustidens Glib Zegdou area during the years 2003 to 2011 by the joint Rasmussen & Simons, 1988, T. andrewsi Matsumoto, 1922, Montpellier/Tlemcen/Oran Universities fieldwork (Tabuce and Titanohyrax nov. sp. (see Rasmussen & Gutiérrez, 2010) et al., 2011). Likewise, T. ultimus, initially described by (Fig. 1). These three species, although smaller than T. ultimus, Matsumoto (1922) based on four heavily worn and isolated are also part of the mammalian megafauna from the Fayum. teeth, has never been rediscovered in the Fayum during the productive joint Duke University/Egyptian Geological From the late early or early middle Eocene of Northwest Museum fieldwork (1977 to present). As for T. angustidens Africa, two other Titanohyrax species are documented, T. and T. andrewsi, only eight and five specimens, respectively, mongereaui Sudre, 1979 from Glib Zegdou, Algeria and T. have been described and recognized according to Rasmussen tantulus Court & Hartenberger, 1992 from Chambi, Tunisia & Simons (1988). Among these rare specimens, Rasmussen & (Fig. 1). These two species were proposed to be the earliest Gutiérrez (2010) recently reattributed, without any comment, representatives of the Titanohyrax lineage, but recently Barrow a maxilla and a mandible of T. angustidens to T. andrewsi, et al. (2010) questioned their generic attribution. These two thereby confusing the systematics of Titanohyrax, a genus with Eocene species differ greatly by their sizes: T. tantulus from an already long and complicated systematic history. Chambi is putatively the smallest Titanohyrax species with a body mass estimated around 23 kg (Tabuce et al., in press), The unexpected recent recognition of a well-preserved whereas T. mongereaui from Glib Zegdou is estimated to have mandible of Titanohyrax andrewsi (MNHN-F-LBE 694) (Figs. weighed about 675 kg (Schwartz et al., 1995), and is thus the 2-3) in the collections of the Muséum National d’Histoire largest African terrestrial mammal from the early Paleogene Naturelle in Paris offers a unique opportunity to better document period. In addition to T. mongereaui, two other very large, as and review a very rare species from the Fayum. Moreover, this 1 PALAEOVERTEBRATA VOL.40 (1)-e4 specimen which was discovered by the French geologist and Andrews (1904b) described but did not figure Megalohyrax paleontologist René Fourtau as early as March-April 1904 ― “minor” based on the maxilla CGM 8818 (figured in Andrews, i.e. two years before the primary description of the two first 1906) now attributed to Megalohyrax eocaenus. He also Titanohyrax specimens by Andrews (1906) ― opens a window described Saghatherium “magnum” (now Geniohyus magnus) on early research in the Fayum. based on the maxilla NHMUK M8398, and “Geniohyus” major (now Bunohyrax major) based on the three associated lower premolars CGM 8980 (figured in Andrews, 1906). At that time, Abbreviations. Andrews referred Geniohyus to the Suidae. AMNH: American Museum of Natural History, New York, Subsequent to this series of short papers, Andrews (1906) USA published additional hyracoid fossils in his comprehensive CGM: Egyptian Geological Museum, Cairo, Egypt ‘Descriptive Catalogue of the Tertiary Vertebrata of the Fayûm, DPC: Duke Lemur Center Division of Fossil Primates, Durham, Egypt’. He figured several specimens already described (see North Carolina, USA above), but also the mandible CGM 8822-3 which he attributed FFZ: Fayum Faunal Zone (Rasmussen et al., 1992) to his small species Megalohyrax “minor”. He also attributed to this species the mandible NHMUK M9220. These two NHMUK: Natural History Museum, Department of Palaeon- specimens were later attributed to the new genus and species tology, London, UK Titanohyrax andrewsi by Matsumoto (1922), CGM 8822-3 MNHN: Muséum National d’Histoire Naturelle, Paris, France being the holotype (Figs. 4-5). SMNS: Staatliches Museum für Naturkunde Stuttgart, Germany After a last expedition in the Fayum in the spring of 1906, YPM: Peabody Museum of Natural History, Yale University, Andrews (1907) figured the mandible NHMUK M9480 New Haven, Connecticut, USA attributed to Saghatherium “magnum” (now the holotype of Thyrohyrax pygmaeus) and he suggested that Geniohyus mirus (NHMUK M9485) may have some relationships with HYRACOIDS FROM THE FAYUM, CHRONOLOGY OF hyracoids instead of suids, a suggestion which is now fully EARLY RESEARCH WITH SPECIAL REFERENCE TO accepted. Among the specimens discovered in 1906, Andrews TITANOHYRAX AND MNHN-F-LBE 694 also cited an enormous canine that he tentatively attributed to the creodont Pterodon africanus. Based on data listed Summary of early research on the NHMUK accession register, this canine is possibly In 1898, the British geologist Hugh J. L. Beadnell from the Egyptian Geological Survey in Cairo, who explored and mapped the eastern and northern border areas of the Fayum Depression, discovered numerous vertebrate remains north of the lake Birket Qarun. In April 1901, he was joined by the pa- leontologist Charles W. Andrews from the British Museum of Natural History for fieldwork in this area. Rapidly,
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