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On Some Ruminant Petrosal Bones and Their Bony Labyrinth from Senèze (Villafranchian, France)

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On Some Ruminant Petrosal Bones and Their Bony Labyrinth from Senèze (Villafranchian, France) Revue de Paléobiologie, Genève (décembre 2018) 37 (2): 443-456 ISSN 0253-6730 On some ruminant petrosal bones and their bony labyrinth from Senèze (Villafranchian, France) Loïc COSTEUR1, Andrea VALLI2, Célia BEAUDOUIN3 & Bastien MENNECART1,4 1 Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001 Basel, Switzerland. E-mail: [email protected] 2 Société Scientifique du Bourbonnais, F-03000 Moulins, France. E-mail: [email protected] 3 Pollen and Cores, 5 rue de l’Eglise, F-68130 Zaessingue, France. E-mail: [email protected] 4 Naturhistorisches Museum Wien, Burgring 7, A-1010 Vienna, Austria. E-mail: [email protected] Abstract We describe the petrosal bone and its associated bony labyrinth (the bony capsule for the inner ear which is the organ of balance and hearing) of five ruminant species of the emblematic lower Pleistocene French locality Senèze. They include two deer (Cervidae) Eucladoceros ctenoides senezensis and Metacervoceros rhenanus philisi and three bovids (Bovidae), the bovine Leptobos (Smertiobos) etruscus, the antilopine Gazellospira torticornis and the caprine Gallogoral meneghinii. We find new morphological characters of both the petrosal bone and the labyrinth that help distinguish between both ruminant families. They relate to the overall shape of the petrosal bone and to specific areas of the bony labyrinth such as the vestibular aqueduct and endolymphatic sac or the lateral semi-circular canal. It confirms previous observations published recently on deer and tragulids (mouse-deer and chevrotains), and adds bovids to our understanding of this structure. This is only a partial study for Senèze since 6 other ruminant taxa are known from there, but it serves as a preliminary work aiming at more often integrating the petrosal bone and bony labyrinth, now easily available using 3D imaging techniques, into taxonomical and phylogenetic studies. Keywords Ruminantia, Cervidae, Bovidae, inner ear, phylogeny, Pleistocene. Résumé Description de pétreux de ruminants et de leurs labyrinthes osseux de Senèze (Villafranchien, France) .- Nous décrivons la portion pétreuse de l’os temporal et le labyrinthe osseux associé de cinq espèces de ruminants de la localité française emblématique du pléistocène inférieur de Senèze. Sont inclus dans cet article deux cerfs (Cervidae) Eucladoceros ctenoides senezensis et Metacervoceros rhenanus philisi ainsi que trois Bovidae: le boviné Leptobos (Smertiobos) etruscus, l’antilopiné Gazellospira torticornis et le capriné Gallogoral meneghinii. Nous mettons en évidence des nouveaux caractères qui permettent de distinguer les deux familles de ruminants étudiées. La forme générale du pétreux et des zones spécifiques du labyrinthe osseux comme l’aqueduc vestibulaire, le sac endolymphatique ou encore le canal semi-circulaire latéral sont particulièrement informatives. Notre étude confirme des observations publiées récemment sur les cerfs et les Tragulidae (chevrotains), et ajoute les Bovidae à notre compréhension croissante de ces structures. Six autres ruminants non étudiés ici sont connus à Senèze, mais cette étude préliminaire contribue à installer le pétreux et le labyrinthe osseux dans les études anatomiques et paléontologiques classiques grâce à l’imagerie 3D qui les rend plus facilement accessibles. Mots-clés Ruminantia, Cervidae, Bovidae, oreille interne, phylogénie, Pléistocène. 1. INTRODUCTION 40Ar/39Ar datings on the abundant volcanic sediments indicate an age bracketed between 2.21 and 2.09 Ma With 40 mammal species, Senèze is one of the richest (Pastre et al., 2015). To date 11 species of ruminants are localities of the European Villafranchian and serves as known from this locality including four deer (Cervidae): a reference fauna for the biochronological timescale Cervalces gallicus (Azzaroli, 1952), Eucladoceros of the European Pleistocene as reference for biozone ctenoides senezensis Heintz, 1970, Metacervoceros MNQ18 (Guérin, 2007). The faunal assemblage comes rhenanus philisi (Schaub, 1941 or M. rhenanus Dubois, from sediments in a volcanic maar setting where often 1904), and Croizetoceros ramosus minor Heintz, 1970, complete specimens in connexion were found. New and seven bovids (Bovidae): Leptobos (Smertiobos) Submitted 20 June 2017, accepted 2 October 2017 Editorial handling: M. Faure DOI: 10.5281/zenodo.2545101 444 L. COSTEUR, A. VALLI, C. BEAUDOUIN & B. MENNECART etruscus (Falconer, 1859), Leptobos (Leptobos) furtivus Theodor, 2010; Orliac et al., 2012, 2017). This is thus Duvernois, 1989, Gallogoral meneghinii (Rütimeyer, only the beginning of a deeper knowledge of ruminant 1878), Gazellospira torticornis (Aymard, 1854), Pro- petrosal bones and bony labyrinths that allows these camptoceros brivatense Schaub, 1923, Pliotragus ardeus phylogenetically relevant structures to be included in (Depéret, 1883), Megalovis latifrons Schaub, 1923, and more comprehensive studies on the evolutionary study Ovis sp. (see up to date faunal list in Delson et al., 2006). of the group. The generic, specific or subspecific status of many of the With the help of non-invasive imaging techniques, which above-mentioned taxa is still discussed, mostly in the have encountered a rapid development in the last 20 years framework of biochronological studies in the Pleistocene enabling the digital reconstruction of otherwise hidden (e.g., see Kostopoulos & Athanassiou, 2005; Croitor, structures, we describe here five petrosal bones and 2006, 2014; Valli & Palombo, 2005 for the specific status bony labyrinths from five fossil ruminant species from of Metacervoceros rhenanus philisi; Duvernois & Guérin, Senèze. This adds morphological data to the description 1989; Duvernois, 1990; Garrido, 2008; Andrés Rodrigo of the deer Eucladoceros ctenoides senezensis and et al., 2010; Masini et al., 2013 for the status of Leptobos Metacervoceros rhenanus philisi and of the bovids etruscus; Crégut-Bonnoure, 2007 or Bellucci & Sardella, Leptobos (Smertiobos) etruscus, Gazellospira torticornis 2015 for the time range of Gazellospira torticornis). and Gallogoral meneghinii. We hope these data will While antlers or horns, teeth, skulls, and postcranial serve for future research in ruminant phylogeny. material have usually been described for ruminants in Senèze (e.g., Schaub, 1941; Guérin, 1965; Heintz, 1970; Duvernois & Guérin, 1989; Duvernois, 1990; Crégut- 2. MATERIAL AND METHODS Bonnoure, 2006, 2007; Crégut-Bonnoure & Guérin, 1996; Delpech & Guérin, 1996), no single petrosal bone Table 1 lists the specimens described in this study with has ever raised interest despite its known relevance scanning information. They all come from the collection in phylogenetic studies (e.g., Webb & Taylor, 1980; of the Natural History Museum Basel (hereafter NMB) O’Leary, 2010). The bony labyrinth, which is embedded and represent only a subset of the ruminant species known within the petrosal bone, is the bony capsule for the inner from Senèze because the petrosal bone was not found for ear, the organ of balance and hearing. It long remained several of them. We CT-scanned the petrosal bones at the vastly unknown for ruminants, both extant and extinct, Biomaterials Science Center in Basel (University Basel) and recent studies have started to show how significant using a Phoenix Nanotom® (General Electric Wunstorf, it could be for phylogenetic analyses (Costeur, 2014; Germany) equipped with a 180 kV / 15 W nanofocus Mennecart & Costeur, 2016a, b; Mennecart et al., 2016; X-ray source. Scanning resolution (Table 1) varies Costeur et al., 2017). With many thousand specimens from 35 µm to 55 µm but this has no influence on the recovered, Senèze has yielded the whole skeleton of final renderings and on measurements taken. The bony many large mammal species and often isolated petrosal labyrinths were segmented using AVIZO 7.0 and AVIZO bones exist for many of them. LITE 9.0 (FEI Visualization Group). Nomenclature Very little comparative data on ruminant, either extant follows O’Leary (2010) for the petrosal bone, and Orliac or extinct, petrosal bones and bony labyrinths have been et al. (2012) and Costeur (2014) for the bony labyrinths. published probably because of the difficulty in accessing The identification of isolated petrosal bones can prove this bone embedded in the cranial cavity and sometimes difficult because of the lack of comparative material and not easily identifiable when found isolated. O’Leary of studies that have described some. The present material (2010) unprecedentedly described detailed and precious was identified thanks to associated identifiable cranial morphological data for some extant and one ruminant specimens like teeth or cranial appendages. Senèze is petrosal bones, but again in a limited sample of 9 taxa, known for the preservation of complete specimens and adding data to Webb & Taylor’s (1980), Guadelli’s the isolated petrosals are often the product of broken (1999), and Mallet & Guadelli’s (2013) samples of skulls that were found in pieces but all in the vicinity recent and fossil taxa. Previous attempts at describing the to each other so that all isolated petrosal bones could be morphology of the bony labyrinth in ruminants occurred attributed to tooth or cranial material, which made their in the late 19th and early 20th century (Hyrtl, 1845; identification easy. Doran, 1878; Gray, 1907, 1908). They are pioneering Nomenclature at higher taxonomic levels such as and seminal works but comparatively contributed little to subfamily or tribe follows Groves & Grubb (2011). our
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    Zitteliana B 32 (2014) 99 The petrosal bone and inner ear of Micromeryx flourensianus (Artiodactyla, Moschidae) and inferred potential for ruminant phylogenetics Loïc Costeur Zitteliana B 32, 99 – 114 München, 31.12.2014 Naturhistorisches Museum Basel, Augustinergasse 2, 4001 Basel, Switzerland, Manuscript received E-mail: [email protected] 26.02.2014; revision accepted 02.06.2014 ISSN 1612 - 4138 Abstract While petrosal bones have a long research history in artiodactyl phylogenetics, the inner ear embedded in this bone has rarely been in- vestigated. I describe here a set of petrosals and the associated inner ears of the Middle Miocene moschid Micromeryx flourensianus from the German locality Steinheim and compare them to the extant musk deer Moschus moschiferus (Moschidae), the four-horned antelope Tetracerus quadricornis (Bovidae) and the white-tailed deer Odocoileus virginianus (Cervidae). Inner ears were reconstructed using high resolution x-ray computed tomography scans. In phylogenenetic reconstructions built on morphological and molecular data, Moschidae has variously been shown to be a sister taxon to Bovidae or Cervidae. Its position hasn’t reached a consensus yet. Studying the inner ear morphology adds new morphological characters that will help resolving this question. Micromeryx flourensianus is an abundant fossil mo- schid and I show indeed that its petrosal bone and inner ear share several similarities with that of the extant musk deer such as a ventral basicapsular groove, a well-developed anterior process of the tegmen tympani, or a fossa for the tensor tympani muscle in the musk-deer that may well have evolved from a Micromeryx-like condition. Inner ears share a thick basal cochlear whorl, a bulky vestibule, or a short and thick cochlear aqueduct.
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