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Specialization for Amphibiosis in Brachyodus Onoideus (Artiodactyla, Hippopotamoidea) from the Early Miocene of France

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Specialization for Amphibiosis in Brachyodus Onoideus (Artiodactyla, Hippopotamoidea) from the Early Miocene of France Swiss J Geosci (2013) 106:265–278 DOI 10.1007/s00015-013-0121-0 Specialization for amphibiosis in Brachyodus onoideus (Artiodactyla, Hippopotamoidea) from the Early Miocene of France Maeva J. Orliac • Pierre-Olivier Antoine • Anne-Lise Charruault • Sophie Hervet • Fre´de´ric Prodeo • Francis Duranthon Received: 28 October 2012 / Accepted: 25 February 2013 / Published online: 16 November 2013 Ó Swiss Geological Society 2013 Abstract A partial cranium of a very large anthracothere strikingly similar to that of extant hippos with: (1) a ventral was unearthed during a palaeontological excavation at basicapsular groove, (2) a sharp crista petrosa, (3) a wide Saint-Antoine-de-Ficalba (Lot-et-Garonne, France; Early prefacial commissure fossa, (4) a reduced mastoid, and (5) Miocene, *18–17.0 Ma). The new material, referred to as an hyperinflated tegmen tympani. Both the disposition of Brachyodus onoideus (Gervais, 1859), documents the cra- the orifices of the head and the petrosal morphology sup- nial features of this species, so far mainly known by dental port a specialization of Brachyodus onoideus to an and postcranial remains. The preserved part of the skull amphibious lifestyle and to potential underwater direc- roughly coincides with the neurocranium and is remarkable tional hearing. for the dorsally-protruding orbits, the importance of the postorbital constriction, the small volume of the braincase, Keywords Anthracotheriidae Á Middle ear Á Burdigalian Á and the gigantic size of the occipital condyle relative to the Underwater directional hearing other elements of the neurocranium. A very careful dis- section of the left auditory region allowed extraction of the left petrosal bone and provides the first description of a 1 Introduction petrosal for Brachyodus. The morphology of the petrosal is The renovation of the RN21 motorway between Ville- neuve-sur-Lot and Agen (SW France) led to the discovery Editorial Handling: D. Becker & D. Marty. by one of us (FP) of a new rich vertebrate locality west to M. J. Orliac (&) Á P.-O. Antoine Á A.-L. Charruault the Saint-Antoine-de-Ficalba village (0.712°E, 44.336°N; Institut des Sciences de l’E´ volution, UMR-CNRS 5554, Fig. 1). The fossiliferous exposure is a thin stri- CC064, Universite´ Montpellier 2, Place Euge`ne-Bataillon, pe, *2.6 km-long (North–South) and 60/110 m-wide 34095 Montpellier, France e-mail: [email protected] (East–West). Fossil-yielding deposits consist of non-indu- rated mica-rich sands and siltstones of fluvio-lacustrine M. J. Orliac origin, known for decades to be late Early Miocene in age Department of African Zoology, Royal Museum for Central (Burdigalian). Fossil remains were found scattered all over Africa, Leuvensesteenweg 13, 30080 Tervuren, Belgium the concerned surface; no significant accumulation, ana- S. Hervet tomical connection, or bone bed was identified during Association Pale´overgne, 24 avenue de l’Abattoir, the excavation undertaken in July 2008. The vertebrate 03450 Ebreuil, France assemblage of Saint-Antoine-de-Ficalba consists primarily F. Prodeo of an abundant chelonian fauna, widely dominated by the Institut National de Recherches Arche´ologiques Pre´ventives, Early Miocene land-dwelling tortoise Testudo promargi- 156 avenue Jean-Jaure`s, 33600 Pessac, France nata von Reinach, 1900 (complete shells), but also including an aquatic/terrestrial testudinid (Ptychogaster F. Duranthon Muse´um d’Histoire Naturelle/AMIS-UMR 5288 CNRS, sp.; connected plates) and a strictly aquatic softshell tur- 35 Alle´es Jules-Guesde, 31000 Toulouse, France tle (Trionyx sp.; connected carapace), both of poor 266 M. J. Orliac et al. biostratigraphical use (Table 1). Among mammals, the concerned stratigraphical interval (*18–17.0 Ma), the rhinocerotids Diaceratherium aurelianense (Nouel, 1866) only anthracothere recorded in Western Europe is and Protaceratherium minutum (Cuvier, 1822), the cervid Brachyodus onoideus (Gervais 1859; Dineur 1981; Gins- Procervulus sp., the moschid Amphitragulus sp., the suid burg 1990; Aguilar et al. 2003; Antunes and Ginsburg Aureliachoerus sp., and the stem cricetid rodent Eucrice- 2003; Lihoreau and Ducrocq 2007). Size and gross anat- todon infralactorensis (Viret, 1930) in Roman and Viret omy of MHNT SAF 001 unambiguously match those of (1930) were notably recognised (Table 1). This assemblage B. onoideus, as illustrated by Dineur (1981). Brachyodus points unquestionably to the ‘base MN3-base MN4’ onoideus is a conspicuous element of the early Orleanian interval, documenting the Early Orleanian European Land European Land Mammal Age (Ginsburg 1990; Bruijn et al. Mammal Age (ELMA), or the Early Burdigalian in terms 1992). Indeed, the first appearance datum of the ‘last an- of standard stages (Mein 1989; Ginsburg 1990; Bruijn et al. thracothere’ of Western Europe is recorded in the locality 1992; Antoine et al. 1997, 2000; Ginsburg and Bulot 2000; of Bru¨ttelen (Switzerland; von Studer 1895; Scherler 2011) Antunes and Ginsburg 2003). dated from the base of the MN3 biozone. B. onoideus is The nature of the fossil-yielding sediments (siltstones very common in localities referred to the middle and upper and sandstones within a clayish matrix), the co-occurrence part of the MN3 biozone (Aguilar et al. 2003; Antunes and of an alligatoroid crocodile (Diplocynodon sp.) with turtles Ginsburg 2003). Its last occurrences are in Artenay (base of strong/strict aquatic affinities (Ptychogaster/Trionyx) MN4; Loire Basin, France; Ginsburg 1990), Nave`re and and a hippo-like rhinocerotid (Diaceratherium aurelian- Marsolan (Fig. 1; base MN4, Garonne Basin, France; Bulot ense; Bentaleb et al. 2006), as well as the taphonomy of the and Ginsburg 1996; Ginsburg and Bulot 2000), Bonrepos- preserved remains (chelonians: complete shells and sur-Aussonnelle (lower MN4, Garonne Basin, France; carapaces but no connected postcranials; mammals: dis- Duranthon 1991; Antoine et al. 1997, 2000), and in the connected elements) point to a fluvial environment, with a ‘‘IVb unit’’ of the Lisbon Miocene Series (18–17.6 Ma, rapid burial of the concerned specimens during a flooding Portugal; Antunes and Ginsburg 2003). episode of a wide river. If dental remains are found all over Europe and relatively A partial cranium of a very large anthracothere, labelled abundant, cranial remains of Brachyodus are very scarce in MHNT SAF 001, was unearthed in Saint-Antoine-de-Fic- contrast (Antunes and Ginsburg 2003). Its cranial anatomy alba during the same field season in 2008. Throughout the was previously described in detail by Dineur (1981), based Fig. 1 Location map of Saint- Antoine-de-Ficalba (Early Miocene, upper MN3) and of selected Early Miocene vertebrate localities from the middle Garonne basin, SW France. Biochronological ages and Brachyodus occurrences are extracted from Baudelot and Collier (1978), Ginsburg (1990), Duranthon (1991), (Bruijn et al. 1992), Antoine et al. (1997, 2000), Ginsburg and Bulot (2000), Bulot et al. (2009) Cranium and petrosal of Brachyodus onoideus 267 Table 1 Vertebrate fauna of Saint-Antoine de Ficalba, SW France dissection of the left auditory region allowed extraction of the (late Early Miocene, Early Orleanian, base MN3-base MN4) left petrosal bone of MHNT SAF 001. Independently, this Supra-familial rank Species Mammalian dissection allowed recognition of another isolated petrosal Neogene zone within the ‘‘isolated petrosal collection’’ from Chilleurs-aux- Bois stored in the NMB collections. Chelonia Ptychogaster sp. Descriptions of petrosals of ‘‘Anthracotheriidae’’ remain Testudo sp. scarce in the literature. Among this paraphyletic family, the Ergilemys sp. latter bones are known so far by an isolated specimen Trionyx sp. referred to as Elomeryx armatus from the White River Crocodylia Diplocynodon sp. Formation, South Dakota (AMNH-VP 579; O’Leary 2010) Serpentes indet. and by two in situ specimens of Bothriogenys sp. from the Mammalia, Diaceratherium (MN3-base MN4) Jebel Qatrani Formation, Fayum, Egypt (DPC 20954, Early Perissodactyla aurelianense Oligocene, *30 Ma; O’Leary et al. 2012) for which only Protaceratherium minutum (MN1-base MN4) the dorsomedial surface has been described. Comparisons Plesiaceratherium sp. (?MN3-MN5) (medium-sized) of the petrosal within hippopotamoids are therefore limited Mammalia, Brachyodus onoideus (MN3-base MN4) to these two taxa and to the living hippos Hippopotamus Artiodactyla amphibius and Choeropsis liberiensis. ?Hyotherium sp. (MN1-MN8) In order to calculate the volume of the tegmen tympani Aureliachoerus sp. (upper MN2-MN5) relative to volume of the petrosal (excluding the mastoid Procervulus sp. (MN3-lower MN5) region) following the protocol of O’Leary et al. (2012), we Amphitragulus sp. (MN1-lower MN4) scanned the extracted left petrosal of MNHT SAF 001 (CTscan Cainotherium sp. (MN1-MN5) facility at the University Montpellier 2, RIO imaging—IBiZA Mammalia, Eucricetodon (MN3) plateform; 1,200 slices, voxel size 35 lm). We reconstructed Rodentia infralactorensis the 3D model of the petrosal using the segmentation tools of The inferred stratigraphical interval, based on mammalian biochro- AVIZO 6.3 (Visualization Sciences Group) and calculated the nology (Ginsburg 1990; Bruijn et al. 1992; Antoine et al. 1997, 2000; corresponding volumes by surface integration. Antunes and Ginsburg 2003), is indicated in terms of Mammal Neogene biozones (MN; e.g., Mein 1989) for each biostratigraphi- cally sound taxon 3 Systematic palaeontology on two partial crania from Chilleurs-aux-Bois, Loiret, Artiodactyla
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