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Re-Evaluation of the Very Large Eomellivora Fricki (Pia, 1939) (Carnivora, Mustelidae, Mellivorinae) from the Late Miocene of Austria

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Re-Evaluation of the Very Large Eomellivora Fricki (Pia, 1939) (Carnivora, Mustelidae, Mellivorinae) from the Late Miocene of Austria Palaeontologia Electronica palaeo-electronica.org Re-evaluation of the very large Eomellivora fricki (Pia, 1939) (Carnivora, Mustelidae, Mellivorinae) from the Late Miocene of Austria Alberto Valenciano, Juan Abella, Ursula B. Göhlich, M. Ángeles Álvarez-Sierra, and Jorge Morales ABSTRACT We re-evaluated the Austrian material for Hadrictis fricki Pia, 1939, from the local- ities Wien XII-Altmannsdorf and Gaiselberg (MN9, Vallesian, Late Miocene), conclud- ing that Hadrictis can be considered as a synonymy of Eomellivora Zdansky, 1924; we therefore named it as Eomellivora fricki. This species is one of the earliest representa- tives of the genus, together with E. piveteaui Ozansoy, 1965. Our phylogenetic analy- ses indicate that Eomellivora forms a monophyletic group, establishing the sister clade of the large and derived Late Miocene Ekorus ekakeran. Eomellivora fricki shows a primitive dental morphology and is the largest species of the genus. This species shows the complexity of the genus Eomellivora, in which large and small species coex- isted since the beginning of the Late Miocene. Alberto Valenciano. Departamento de Geología Sedimentaria y Cambio Medioambiental. Instituto de Geociencias (CSIC, UCM). Calle José Antonio Novais 12, 28040. Madrid, Spain and Departamento de Paleontología Universidad Complutense de Madrid (UCM), Facultad de Ciencias Geológicas UCM. Calle José Antonio Novais 12, 28040, Madrid, Spain. [email protected] Juan Abella. Universidad Estatal Península de Santa Elena. Kilómetro 1,5 de la Vía Santa Elena- La Libertad. Edificio Instituto de Investigacion Científica y Desarrollo Tecnológico INCYT), Segundo Piso. 240210 La Libertad, Ecuador. [email protected] and Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona. Edifici Institut de Ciència i Tecnologia Ambientals - Institut Català de Paleontologia (ICTA-ICP), Campus de la UAB, Carrer de les Columnes s/n, 08193 Cerdanyola del Vallès, 08193, Barcelona, Spain. [email protected] Ursula B. Göhlich. Naturhistorisches Museum Wien, Dept. Geology and Paleontology Burgring 7, 1010 Vienna, Austria. [email protected] M. Ángeles Álvarez-Sierra. Departamento de Geología Sedimentaria y Cambio Medioambiental. Instituto de Geociencias (CSIC, UCM). Calle José Antonio Novais 12, 28040. Madrid, Spain and Departamento de Paleontología Universidad Complutense de Madrid (UCM), Facultad de Ciencias Geológicas UCM. Calle José Antonio Novais 12, 28040, Madrid, Spain. [email protected] Jorge Morales. Departamento de Paleobiología. Museo Nacional de Ciencias Naturales-CSIC, Calle José Gutiérrez Abascal, 2, 28006, Madrid, Spain. [email protected] Valenciano, Alberto, Abella, Juan, Göhlich, Ursula B., Álvarez-Sierra, M. Ángeles, and Morales, Jorge, 2017. Re-evaluation of the very large Eomellivora fricki (Pia, 1939) (Carnivora, Mustelidae, Mellivorinae) from the Late Miocene of Austria. Palaeontologia Electronica 20.1.17A: 1-22 palaeo-electronica.org/content/2017/1830-the-large-eomellivora-fricki Copyright: © April 2017 Palaeontological Association VALENCIANO ET AL.: THE LARGE EOMELLIVORA FRICKI Keywords: Eomellivora; Hadrictis; Mellivorinae; Mustelidae; Miocene Submission: 15 June 2016 Acceptance: 8 April 2017 INTRODUCTION to Mellivorinae in this manuscript), Eomellivora Zdansky, 1924, Erokomellivora Werdelin, 2003, Mustelidae is the most diverse family within Hadrictis Pia, 1939, Howellictis Bonis et al., 2009, the currently existing Carnivora, comprising 57 Mellivora Storr, 1780 and Promellivora Pilgrim, extant species of weasels, martens, polecats, bad- 1932. Some of them, such as Ekorus and Eomelliv- gers and otters (Larivière and Jennings, 2009). ora (=Hadrictis), reached a significantly large size, The Mellivorinae Gray, 1865, is the subfamily con- which is why they are considered as giant muste- taining Mellivora capensis (Schreber, 1776) as the lids. Eomellivora and Ekorus were one of the larg- only living representative, commonly known as the est and most hypercarnivorous mustelids ever ratel, or honey badger. It is the largest African ter- known; they were larger than the extant wolverine, restrial mustelid, weighing between 6.2 and 13.6 Gulo gulo, but according to their cranium dimen- kg, and with a distribution range from Africa to sions, they were smaller than the Oligobuninae India (Larivière and Jennings, 2009). It is a gener- Megalictis ferox (Werdelin, 2003; Valenciano et al., alist and opportunistic predator, with a wide range 2015, 2016). Eomellivora has been described in of prey; it presents clear regional differences in its Asia, North America, Europe and Africa (e.g., Wol- diet, which mostly involves rodents, other carniv- san and Semenov 1996; Morales and Pickford, orans (e.g., Suricata, Felis, Cynictis, Ictonyx), 2005; Valenciano et al., 2015), spanning from the amphibians, reptiles, birds and invertebrates, but Middle (MN8) to the Late Miocene (MN13). The also includes roots, berries and fruits (Begg et al., first complete review of the genus was conducted 2003; Larivière and Jennings, 2009). Notwithstand- by Wolsan and Semenov (1996); it concluded that ing the fact that only one monospecific genus has this genus represents a single lineage of E. wimani survived up to the present, this subfamily was more Zdansky, 1924, which they subdivided into the diversified in the past. This mellivorine diversity is chrono-subspecies E. wimani piveteaui for the well reflected in the Neogene fossil record, Vallesian (MN9-10) specimens, and E. wimani although the remains are mostly fragmentary wimani for the Turolian/Ventian (MN11-13) ones. (Baskin, 1998; Morales et al., 2015, Werdelin and More recently, a sample of E. piveteaui Ozansoy, Peigné, 2010). The origin of the subfamily is 1965, from Batallones (Late Miocene, MN10, unclear, but most likely arose in Eurasia or Africa, Madrid, Spain) has been described by Valenciano during the Middle Miocene and early Late Miocene et al. (2015); this has enabled E. piveteaui, E. with Sivamellivora Kretzoi, 1942, Mellalictis Gins- wimani, E. ursogulo (Orlov, 1948) and E. hungarica burg, 1977 and Eomellivora ?tugenensis Morales Kretzoi, 1942, to be accepted as valid species. and Pickford, 2005. Sivamellivora necrophila (Pil- However, neither Wolsan and Semenov (1996) nor grim, 1932) comes from the Lower Siwaliks (India), Valenciano et al. (2015) analyzed the very large- Chinji formation ca. 14-11.2 m.y.a. (Patnaik, 2013) sized mustelid Hadrictis fricki Pia, 1939. and is based on very few teeth and an edentulous Hadrictis fricki was described by Pia (1939) mandible. Mellalictis mellalensis Ginsburg, 1977 from the Austrian locality of Wien XII-Altmannsdorf, from Beni Mellal (Morocco) ca. 12.5-11.2 m.y.a. Late Miocene (MN9), by a largely fragmented skull (Werdelin and Peigné, 2010) is known by its iso- that includes a worn P4-M1. Subsequently, Zapfe lated teeth and postcranial fragments. It is uncer- (1948) described a hemimandible with a worn den- tain, however, whether it is a Mellivorinae or a tition as H. fricki from Gaiselberg (Austria), Late mustelinae (Bonis et al., 2009). Eomellivora Miocene (MN9). The systematic position of Hadric- ?tugenensis Morales and Pickford, 2005 from the tis is doubtful - it constitutes a valid genus for some Ngorora formation (Kenya), ca. 12 m.y.a., is a authors (Pia, 1939; Zapfe, 1948; Kretzoi 1942, poorly known mustelid of medium size that could 1965; Ozansoy, 1965; Webb, 1969; Ginsburg, constitute an ancestral form of Eomellivora (Valen- 1977), whereas for others it is a synonym of ciano et al., 2015). Eomellivora (Werdelin, 1996, 2003; Peigné et al., Since the beginning of the Late Miocene, new 2006). However, the mandible of H. fricki described genera of mellivorines radiated throughout Eurasia by Zapfe (1948) has been ignored and never com- and Africa, e.g., Ekorus Werdelin, 2003 (assigned pared with any other species of Eomellivora. The 2 PALAEO-ELECTRONICA.ORG FIGURE 1. Vienna Basin with geographic and stratigraphic position of localities (asterisks) (modified after Har- zhauser et al., 2004, figures 1 and 2). objectives of the present manuscript are twofold: stage the foregone disintegration of the Central (1) to re-describe and re-measure the material Paratethys has restricted Lake Pannon to the Pan- described by Pia (1939) and Zapfe (1948), and (2) nonian Basin system. The first brackish-lacustrine to re-evaluate the systematic position of H. fricki in and subsequent fluvial-influenced deposits of Lake the light of the new evolutionary framework of Pannon are biostratigraphically subdivided into the Eomellivora proposed by Valenciano et al. (2015). Pannonian A to H biozones, based on molluscs (Papp, 1951). Both sites yielded a Late Miocene Localities and Geological Setting (Pannonian) vertebrate fauna (e.g., Pia, 1939; The- Both Austrian localities, Wien XII-Altmanns- nius, 1948; Zapfe, 1949) including fossils of the dorf and Gaiselberg, are situated in the Vienna three-toed horse Hippotherium sp., which FAD Basin, which forms the northwestern part of the (First Appearance Datum) is MN9. Pannonian Basin (Figure 1). During the Pannonian 3 VALENCIANO ET AL.: THE LARGE EOMELLIVORA FRICKI TABLE 1. Fossil vertebrate fauna Wien XII-Altmannsdorf TABLE 2. Fossil vertebrate fauna from Gaiselberg (Griesergasse, Oswaldgasse) (updated by U.B.Göhlich, (uddated by U.B. Göhlich, based on Zapfe (1949), The- based on Pia (1939) and Thenius (1948)). nius (1950), Pickford (2016) and J. Giaourtsakis (per- sonal commun., Athens, 2014). Wien XII-Altmannsdorf
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