Complete description of the skull and mandible of the giant mustelid Eomellivora piveteaui Ozansoy, 1965 (Mammalia, Carnivora, Mustelidae), from Batallones (MN10), late Miocene (Madrid, Spain)

ABSTRACT We describe cranial, mandibular, and dental remains of five individuals of the giant mustelid Eomellivora piveteaui Ozansoy, 1965, from the late Miocene (MN10) site of Cerro de los Batallones (Madrid, Spain)—the first complete cranial remains recorded for this species and the most complete remains of the genus. This new sample enables a review of the systematic status of Eomellivora, leading us to accept as valid the species E. piveteaui Ozansoy, 1965, E. wimani Zdansky, 1924, E. ursogulo (Orlov, 1948), and E. hungarica Kretzoi, 1942. Our phylogenetic hypothesis indicates that Eomellivora is the sister taxon of the extant Mellivora capensis and E. piveteaui had a common ancestor within the crown group E. wimani–E. ursogulo. Eomellivora piveteaui was specialized for a more hypercarnivorous diet than the largest extant terrestrial mustelids, although it also had some derived bone-crushing adaptations. Eomellivora piveteaui had an active predatory role in the late Miocene carnivore faunas, exploiting both small and relatively large prey. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP


INTRODUCTION
The Cerro de los Batallones fossil sites (late Miocene [MN10]) have yielded some of the most interesting, richest, and best-preserved Neogene mammal assemblages of the Iberian Peninsula . It is a system of nine distinct sites located on a structural butte in the south of the province of Madrid, Spain (Fig. 1). Fossil remains indicate a late Vallesian age (ca. 9 Ma), for all the sites. However, there are slight differences in composition in micro-and macromammals among the different fossil deposits that have been attributed to minor temporal differences indicating, for instance, that Batallones-10 is older than Batallones-3 (L opez- Antoñanzas et al., 2010;Siliceo et al., 2014). All these Batallones sites were cavities that acted as natural traps for vertebrates. The fossiliferous deposits are embedded in marls and are composed of two types of assemblages: (1) subsurface cavities (e.g., Batallones-1 and Batallones-3) interpreted as carnivore traps and (2) doline-like depressions (e.g., Batallones-4 and Batallones-10) interpreted as herbivore traps. The two types differ in their stratigraphic relative position, internal stratigraphic architecture, taxonomic composition, and taphonomic features (Pozo et al., 2004Abella, 2011;Abella et al., 2011;Domingo et al., 2011Domingo et al., , 2013Calvo et al., 2013).
The numerous excavation campaigns in the different Batallones localities have yielded a rich assemblage of vertebrate fossils, including fishes, amphibians, reptiles (small and giant tortoises, lizards), and predatory birds P erez-Garc ıa and Murelaga, 2013), as well as micromammals (L opez-Antoñanzas et. al., 2010), herbivores S anchez et al., 2009R ıos et al., 2013), and carnivorans, with the latter constituting the most diverse sample (e.g., Ant on et al., 2004aAnt on et al., , 2004bPeign e et al., 2005Peign e et al., , 2008Salesa et al., 2006Salesa et al., , 2008Salesa et al., , 2010Salesa et al., , 2012Abella, 2011;Abella et al., 2011Abella et al., , 2013Valenciano et al., 2012Valenciano et al., , 2013. The giant mustelid was identified from Batallones-3 by Valenciano et al. (2012) as Eomellivora piveteaui Ozansoy, 1965. The richness of this sample is remarkable in that it includes relatively complete cranial and postcranial material, representing the most complete remains known of the genus. The craniodental specimens belong to at least five individuals and include two skulls, eight hemimandibles, and several isolated teeth (Figs. 2,3). Eomellivora played an important role in the late Miocene Hipparion faunas, being one of the largest and most hypercarnivorous mustelids ever known, larger and more hypercarnivorous than the extant wolverine Gulo gulo (Valenciano et al., 2013). Eight species of Eomellivora have been described from 18 different localities in the Old and New World (e.g., Tobien, 1955;Lupt ak, 1995;Wolsan and Semenov 1996;Morlo, 1997;Morales and Pickford, 2005;Koufos, 2012), spanning the middle (MN8) to late (MN13) Miocene. However, Wolsan and Semenov (1996) considered only a single valid species (E. wimani Zdansky, 1924), which they subdivided into two chronosubspecies: the Vallesian (MN9-10) E. w. piveteaui and the Turolian/Ventian (MN11-13) E. w. wimani. The aim of the present paper is to describe in detail the morphology of the complete skull, mandible, and dentition of Eomellivora from Batallones (including a three-dimensional [3D] digital model that removes some of the taphonomic distortion) and to study the systematic position of Eomellivora and its included taxa.

Three-Dimensional Model
The skull of E. piveteaui from BAT-3 is almost complete, but crushed, due to the compaction of the sediments resulting in both fragmentation and deformation. This kind of distortion has been previously reported in the carnivoran cranial material from BAT-1 (Domingo et al., 2013). Although the exceptional preservation of the skull permits reliable observations of morphological details, the distortion makes it difficult or impossible to accurately interpret three-dimensional (3D) aspects (Ant on et al., 2004a). The recovery of the original shape through 3D reconstruction of individuals with distortions improves overall morphological descriptions and constitutes a necessary first step toward a correct anatomical reconstruction. Two main deformation forces in BAT-3'09.1000 on the sagittal and coronal planes have been detected through observation of the relative position of symmetric structures. A surface 3D model has been obtained using a Next Engine 2020i scanner (Kuzminsky and Gardiner, 2012). In order to restore the original morphology, a shear deformation was applied to the sagittal plane of the 3D model through the CAD software Autodesk 3D Studio Max, with an angle of shearing of 18.3 estimated. Similarly, a 3.4 shearing deformation has been applied to the coronal plane. Both shearing amounts have been estimated based on realignment of paired structures. Although observed, the quantification of the minor dorsoventral distortion is difficult to estimate, thus preventing its correction. The corrected interactive 3D model has been created in PDF 3D format (which can be opened and manipulated in standard PDF viewers) for easy accessibility and comparison (Supplementary Data 1, Figs. S1, S2).
Differential Diagnosis-Eomellivora piveteaui differs from E. wimani, E. ursogulo, and E. hungarica in its longer muzzle, more sectorial dentition, and reduced p4 mesial accessory cuspid. Additionally, it differs from E. wimani and E. ursogulo in: more slender P2, P4, M1, p2, m2; P2 with a weak lingual cingulum; lingual platform of M1 not enclosed completely the protocone; and p3 without a buccolingual platform. Differs from E. wimani in much more slender P2 and p2; less concavity in the buccal wall of P3-P4; P3 with mesial accessory cusp less developed; M1 with more substantial metacone and slender protocone not centrally located; and slender m1 talonid with sharper hypoconid. Differs from E. ursogulo in: P3 without two distal accessory cusps; P4 with mesodistally reduced protocone and without additional cusp in the parastyle; lingual platform of M1 without inflexion; p2 without distal accessory cuspid; p3 mesial accessory cuspid not developed and with a much more slender distal area; and more primitive m2 with three sharper cuspids in a central position along the mesodistal axis. Differs from E. hungarica in: much longer mandible without reduction of the dental series; p4 shorter and more slender, with more vertical main cuspid; and m1 more slender and with no lingual cristid.

Description
Skull and Upper Dentition-Two skulls (BAT-3'09.1000 and BAT-3'13.185) of Eomellivora have been found at Batallones. BAT-3'09.1000 ( Fig. 2A1-4) has an approximate basal cranial length of 182.52 mm. Its preservation is relatively good, despite the fact that it is compressed and warped approximately 18.3 in the sagittal plane and 3.4 in the coronal plane (Fig. 2). The nuchal and ventral regions are also quite damaged. BAT-3'13.185 (Fig. 2B1, 2) is lateromedially compressed, and the nuchal and ventral regions are quite damaged. The foramen magnum, occipital condyles, and basisphenoid and basioccipital bones are missing in both specimens. The dentition of BAT-3'09.1000 is partially dissolved by soil acids, and also partially broken, with some of the cuspids of the lower teeth being impacted into the upper dentition. The dentition of BAT-3'13.185 is relatively good and retains P2-M1.
3'13.185. On the maxillary bones, there is a moderately deep and rounded fossa in the rostral margin of the orbit that extends dorsally. The large infraorbital foramen (6.58 mm width and 12.83 mm height in BAT-3'13.185) is located under the frontal process of the maxilla and above the mesial edge of the P4 paracone. The rostral margin of the orbit ends at the level of the distal margin of the P4 paracone. BAT-3'09.1000 has a weak temporal crest and a well-developed sagittal crest that extends caudally towards the external occipital protuberance, where it divides into the nuchal crests, forming a 'Y' pattern. The external occipital protuberance does not exceed the nuchal region. In caudal view (Fig. 2A4), the nuchal area is triangular, rather flat, and has a large area of insertion on the supraoccipital bone.
The zygomatic arches are robust in their rostral and caudal areas and are especially robust near the glenoid cavity. Both M. masseter pars superficialis and M. masseter pars profunda have their origin on the ventrolateral side of zygomatic arch. The frontal processes of the zygomatic arches are triangular, high, and rostrocaudally broad.
Ventrally, the tooth rows are rectilinear between C1 and P4. A diastema separates I3-C1. Neither the pterygoid region nor the hamulus pterygoideus processes are well preserved. The tympanic bulla is partially inflated, with a rugose ventral surface and a large and rounded external auditory meatus. The ventral parts of the nuchal crest in conjunction with the mastoid process are well developed and are rostrolaterally widened. The nuchal crest is concave and projects laterally, which creates large attachment areas for M. zygomatic temporalis on the dorsal side and M. obliquus capiti cranialis on the caudal side. The mastoid process is robust and is situated lateral to the tympanic bulla. The paroccipital process is caudal to the auditory bulla. BAT-3'09.1000 possesses a medium-sized paroccipital process, narrow, triangular in section, and projecting laterocaudally. The M. digastricus originates on the lateral side of the process.
The preserved upper dentition includes I3, C1, P2, P3, P4, and M1. The I3 BAT-3'09.688 ( Fig. 2C1-2) is caniniform, with a single cusp curved distally. On the lingual side, there is a sharp crista as well as a marked cingulum. It has a massive root that is oval in cross-section. The C1 BAT-3'08.635 (Fig. 2D) has a long FIGURE 4. Relationships between lengths (L) and widths (W) of the upper dentition in Eomellivora. Sources: SH (Shangyingou), Zdansky (1924); LI (Liuwangou), Zdansky (1924); GRT (Gritsev), Wolsan and Semenov (1996); NO (Novaya Emetovka), Orlov (1948); GY € O (Gy€ orszentm arton), Kretzoi (1965), KRF (Kern River Formation site 50), Stock and Hall (1933); CIM (Cimislia), Wolsan and Semenov (1996); YAS (Yassi€ oren), Ozansoy (1965), and for P4 and M1, estimations based on pictures of MNHN-TRQ-1005, instead of the evidently confused original data provided in Ozansoy (1965); WSS (Wissberg), Tobien (1955); RPI (Ravin de la Pluie), Koufos (2012); BAT (Batallones), this paper; LVF (Los Valles de Fuentidueña), Crusafont-Pair o and Ginsburg (1973), and for p2, this paper; KLF (Kalfa), Lungu (1978), and for M1, this paper, estimations based on text- fig. 9 in Lungu (1978), instead of the apparently mistaken original data given in table 2; GRE (Grebeniki), Orlov (1948); NG (2/11 locality, Ngorora Formation), Morales and Pickford (2005); BOJ (Borsk y Sv€ at y Jur), Lupt ak (1995); CSA (Cs arkvar), Kretzoi (1942); POL (Polg ardi 2). crown, wide at the base. The mesial and distal cristae are corroded. The crown has a small distal cingulum, and the root is robust and buccolingually wide near the cementoenamel junction. BAT-3'13.185 has an alveolus for P1 (Fig. 2B2); the P2 is two rooted and has a high and mesially oriented single cusp (Fig. 2B1, E1, 2). It is buccolingually rotated relative to the tooth row. The mesial and distal cristae have sharp edges. It widens distally and has a marked distolingual cingulum. The P3 (Fig. 2B2) is triangular in occlusal view, bicuspid, and two rooted. Its cusps are distolingually oriented. The main cusp is very high, and the distal accessory cusp is not very prominent. There is a marked lingual expansion, a broad lingual cingulum, and a concavity in the buccal wall. The P4 of BAT-3'13.185 (Fig. 2B2) is triangular, and robust. It has a very small and low parastyle located on its mesial cingulum (Fig. 2B1). The protocone is subconical, robust, and located in line with the parastyle. There is a slight mesial inflection between the paracone and the parastyle. There is a poorly developed concavity in the buccal wall. The paracone and metastyle form an elongate blade. The paracone is the highest and largest cusp, occupying over half of the total length of the tooth. It is triangular and distally inclined. The distal crista of the paracone has a somewhat concave slope that fuses to the mesial crista of the metastyle via a 'U'-shaped valley. The M1 (Fig. 2B2) is rectangular, with the buccal wall narrower than the lingual one. The paracone is conical and situated on the mesiobuccal corner. It is the highest and most robust cusp, although the metacone is also quite well developed. There is a markedly swollen buccal cingulum. The stylar area of the M1 is enlarged. The protocone is robust, subconical, and mesiolingually located. There is a very swollen lingual platform that does not completely enclose the protocone.
Mandible and Lower Dentition-The mandible of the Batallones Eomellivora is long and robust ( Fig. 3A1-3, B1-2, C). The specimens BAT-3'08.526 (Fig. 3A1-3) and BAT-3'09.1000 (Fig. 3C) have a total length of 120.79 and 134.16 mm, respectively. The tooth row is straight and aligned with the articular process. The mandibular corpus is deep, approximately twice the height of the m1, and is mediolaterally widened at the level of the paraconid of m1. The ventral margin is convex at the level of the ascending ramus and the mandibular symphysis, but it is concave from the p3 to the trigonid of m1. On the lateral side, there are two rounded mental foramina, one beneath p2 and the other beneath p3. The mandibular symphysis is rather vertical, being more evident in the larger hemimandibles (Fig. 3C). The muscle attachment of M. digastricus is on the caudoventral edge of the mandibular corpus and reaches rostrally to beneath the m1 trigonid.
The ascending ramus is rostrocaudally broad. Its dorsal border is sharp and vertically oriented. The coronoid process is laterally rotated with an angle of »75 with respect to the articular process. The M. temporalis pars profunda and the M. temporalis pars superficialis have their areas of insertion on the medial and lateral sides of the ascending ramus, respectively. The masseteric fossa is large, oval, and relatively deep. Its rostral margin lies at the level of the m2. Inside the masseteric fossa, there is a noticeable and deep insertion area for the M. zygomaticomandibularis. The M. masseter pars superficialis inserted on the ventrolateral edge of the angular process, which also has a large insertion for the M. masseter pars profunda near its ventrolateral margin. The articular process is large and is widely separated from the angular process. The angular process is robust, caudally directed, and on its medial side has a well-developed muscular attachment for M. pterygoideus medialis.
There are two deciduous teeth (dp3 and dp4) associated with BAT-10'12-G2-4 ( Fig. 3D1-8). The dp3 (Fig. 3D3-5) is sigmoidal, two rooted, and tricuspidate. The mesial accessory cuspid is very low; there is a small notch between it and the main cusp. The main cuspid is high, lingually widened at the base and somewhat mesially placed, with two sharp cristids. The distal accessory cuspid is located on a cingulid. It is the lowest cuspid and has a large valley between it and the main cuspid. The dp4 ( Fig. 3D6-8) is fragmentary. It is also two rooted, elongated, and much larger than dp3. It preserves the buccal wall of the paraconid, a complete protoconid, and a small fragment of the hypoconid with a tiny linguocclusal wall. The trigonid is similar to that of the m1; it has no metaconid and possesses a convex buccal wall. The paraconid is low and reduced relative to the protoconid. The protoconid is the highest cuspid. There is a wide valley between the trigonid and the talonid. The adult lower dentition includes the c1, p2-p4, and m1-m2. All the cuspids are aligned on the longitudinal axis. No lower incisors are preserved. The lower canine of BAT-3'08.526 (Fig. 3A 1-3) is robust and conical and has heavy occlusal wear. It has an oval cross-section, slightly  broader at the base and curved distally. Some longitudinal lines radiate from the apex to the crown. The p1 is not preserved, but there is a single small alveolus in the larger mandibles. The p2 (Fig. 3A1-3, C) is two rooted, with elliptical shape formed by one cuspid mesially displaced. It is buccolingually rotated relative to the tooth row and is elongated but with a slight distal widening. It has small mesiobuccal and noticeable distobuccal cingulids. The p3 (Fig. 3A1-3) is biscuspidate. It has a pronounced buccolingual widening. The main cuspid is elongated and buccolingually compressed; its mesial and distal cristids have sharp edges. The distal accessory cuspid is low and has a blunter morphology. The mesial, buccal and lingual cingulids are well developed. The p4 (Fig. 3A1-3, B1-3, C) has three cuspids and is subrectangular with a slight distal broadening. It is a relatively long tooth compared with the total length of m1. It has a very small mesial accessory cuspid. Its main cuspid is buccolingually compressed; the crown of p4 is higher than that of the m1 paraconid. Its mesial and distal cristids have sharp edges. There is a noticeable intercuspid notch between the main cuspid and the distal accessory cuspid. The distal accessory cuspid is high, and  Kaakinen et al. (2013); and Vangengeim and Tesakov (2013); B, calibrated phylogenetic hypothesis based on Appendices 1-2 (see Fig. 9 for more details). blunt in BAT-3'09.1000, and low in BAT-3'13.230, suggesting some degree of variability in this feature. Its main cuspid is inclined distally toward the m1 (see Appendix 1). There is an occlusally projected mesial cingulid and a larger cingulid surrounding the distal accessory cuspid. The m1 (Fig. 3A1-3, B1-3, D1, 2) is elongated and with a buccolingually compressed talonid. The trigonid is a trenchant blade that occupies almost three-fourths of the total length of the tooth and has a prominent FIGURE 8. Ratio diagram of all Eomellivora described in the bibliography. The dental measurements of Gulo gulo B.1280 serve as the standard for comparison (solid line at Y axis equal to 100). Legend and sources, the same as Figure 4. carnassial notch. The protoconid is higher than the paraconid, and the two cuspids are located in the same plane. There is no metaconid, but there is a cristid that reaches the lingual area of the talonid enclosing a vestigial basin. The talonid is high and located in line with the trigonid cuspids; it is composed of a buccolingually compressed hypoconid occupying most of the talonid and a small centrally located hypoconulid only perceptible in the not fully erupted and unworn m1 of BAT-10'12-G2-4 ( Fig. 3D1, 2). There is no entoconid. The talonid has noticeable buccal and lingual cingulids. The m2 (Fig. 3D1, 2) is uniradiculate, tricuspidate, and oval in occlusal outline. As in the m1, the trigonid measures three-fourths of the total length; its cuspids are located in the same plane as the talonid cuspids of the m1. It has a low and sharp paraconid, situated on a cingulid. The protoconid is the highest and sharpest cuspid; it is located in a central position. There is a small notch between the paraconid and the protoconid. The talonid has a central hypoconid on a well-developed cingulid.

Systematic Discussion
The taxonomy of the genus Eomellivora has remained problematic because a substantial number of species have been named based on very scarce remains and the range of dental variation is largely unknown. For this reason, we believe that the completeness of the sample of Eomellivora from Batallones (Figs. 4-9) presents a great opportunity to shed light on the systematics of the genus (Table 3).
Eomellivora piveteaui Ozansoy, 1965, was first described on the basis of a partial right hemimandible with i2-p4 and m1 trigonid, a fragment of maxilla with P4 and M1, and an isolated M1 from Yassi€ oren (Turkey), MN9 (Fig. 6A1-3). Wolsan and Semenov (1996) included this taxon in Eomellivora wimani at the subspecific rank of E. wimani piveteaui, and they chose the Yassi€ oren hemimandible MNHN-TRQ-1004 as lectotype (Table 3). The similarities between the material from Batallones and the Yassi€ oren Eomellivora are striking (Valenciano et al., 2012), especially BAT-3'09.1000, which has dentition of similar size and morphology  to the material described by Ozansoy (1965). However, there are some differences between the material, such as the smaller relative size of the p2 and M1 and the more robust p3 in the Batallones-3 specimens (Fig. 8).
The Eomellivora from Batallones includes five individuals (four from Batallones-3 and one from Batallones-10) and shows great size variation (Figs. 4,5,8,Tables 1,2). This variability reaches up to 20% of the tooth length (e.g., m1) and appears to indicate the existence of marked sexual dimorphism in the genus, as occurs in other extant and extinct mustelids (e.g., Zakrzewski, 1967;Moors, 1980;Hunt and Skolnick, 1996;Baskin, 1998;Ewer, 1998;Larivi ere and Jennings, 2009). Thus, we believe that sexual dimorphism can help us to interpret the systematics of Eomellivora, and explain the larger size of some Batallones specimens in relation to the material from Turkey. The two samples are similar in the absence of a p2 distal accessory cuspid, and slight mesial accessory cuspid in p3; similar robustness in p4 and P4; P4 shows a similar concavity in its buccal wall, with a robust protocone in line with the parastyle; and the M1 has a conical paracone and a well-developed metacone, with a robust, subconical, and mesiolingually located protocone as well as buccal and lingual cingula. Therefore, the Eomellivora from Batallones-3 is conspecific with E. piveteaui from Yassi€ oren (Fig. 7). The specimen of Eomellivora from Batallones-10 cannot be completely compared with the Yassi€ oren sample because the only comparable element, the m1 trigonid, is insufficient for comparative assessment. However, this specimen is metrically and morphologically similar to the Batallones-3 specimens; therefore, we are confident in identifying it as E. piveteaui. An isolated m1 without roots belonging to a young adult described as Mellivorine, gen. et sp. indet., from the Wissberg locality (Germany), MN9 (Tobien, 1955), was considered to belong to E. wimani piveteaui by Morlo (1997). This m1 is similar in size to the largest teeth of the E. piveteaui sample from Batallones-3, and the morphology of the m1 talonid is similar to the m1 from Batallones-10; thus, we assigned it to E. piveteaui (Fig. 7, Table 3).   Lungu (1978) ascribed a complete dental sample of one individual from Kalfa (Moldova), MN9, to E. piveteaui. With the exception of Wolsan and Semenov (1996) paper, no other papers seem to have taken this description into account. This material is comparable to the lectotype of E. piveteaui from Yassi€ oren, except for the ascribed m2 (UST-CLF-N1-2027) that actually belongs to a hyaenid. The dentition from Kalfa is similar to that of E. piveteaui from Batallones, only differing in the more developed M1 distal platform and a larger length of M1, p3, and p4, and more robust M1 and m1 in the Eomellivora from . Due to the larger size of the E. piveteaui from Kalfa and the older age of this locality in relation to Eomellivora from Batallones, these differences can be explained in terms of temporal differences or sexual dimorphism, and we agree with the taxonomic determination of Lungu (1978) (Table 3). Crusafont-Pair o and Ginsburg (1973) described very fragmentary material of the taxon Eomellivora liguritor, from the late Miocene (MN9) locality of Los Valles de Fuentidueña (Spain), consisting of an isolated M1 and a P4 without its protocone (Ginsburg et al., 1981). This species was synonymized with E. wimani piveteaui by Wolsan and Semenov (1996). The dentition is similar in size to E. piveteaui from Kalfa and E. wimani from China, but based on the remains of Kalfa, we think this Spanish material should be assigned to E. piveteaui (Table 3). Koufos (2012) described and classified an isolated and worn M1 from Ravin de la Pluie, Greece, MN10, as E. wimani. However, it is metrically and morphologically similar to the M1 from Kalfa; thus, we reclassified it as belonging to E. piveteaui (Table 3).
Perunium ursogulo from Grebeniki (Ukraine) was described by Orlov (1948), based on a fragment of skull and two mandibles (Fig. 6C1-3). The age of this locality is MN11 (Wolsan and Semenov, 1996), although recent studies suggest an greater age (MN10; Vangengeim and Tesakov, 2013). Due to the strong similarity with Eomellivora, Ozansoy (1965) subsumed Perunium in this genus and Wolsan and Semenov (1996) synonymized the species with E. wimani wimani. The dental morphology of the Ukrainian sample is intermediate between E. piveteaui and E. wimani, but it possesses significant differences from both species (Fig. 6). On the one hand, it shares a poorly developed buccal concavity in P4, an M1 metacone that is more developed, and a mesolingually positioned M1 protocone with E. piveteaui. On the other hand, it differs from E. piveteaui in: relatively shorter muzzle; more robust dentition (especially the p3 with a very strong distal widening); a mesiodistally reduced P4 protocone; more accessory cuspids in p2-p4; and larger and more derived m2, with a reduced single cuspid placed in a central position, together with a mesiodistal cristid. Moreover, E. ursogulo shares with E. wimani, robust P2, P3, p2, and p3; strongly developed lingual cingulum in P2 and mesial cingulum in P3; more highly developed premolar accessory cuspid; and the lingual platform of M1 completely encloses the protocone. Although E. ursogulo seems to possess a mixture of morphological features of E. piveteaui and E. wimani, it also appears to be a highly derived species, having some notable autapomorphies that, in our opinion, support the taxon as a separate species (Table 3). The autapomorphies include: small additional cusp in the P4 parastyle; P3 with two distal accessory cusps; lingual platform of M1 with an inflexion; and highly developed p3 mesial accessory cuspid. Eomellivora ursogulo has also been diagnosed outside Ukraine from Borsk y Sv€ at y Jur (Slovakia), MN11 (Lupt ak, 1995). However, a more recent paper by Kov a c et al. (2006) revised the age of this site to MN9 and identified the material as E. wimani. Due to the scarcity of the material, we prefer to classify the Slovakian specimens as Eomellivora sp. (Fig. 7, Table 3).
Eomellivora hungarica Kretzoi, 1942, from Polg ardi 2 (Hungary), synonymized as E. wimani wimani (Wolsan and Semenov, 1996), was described on the basis of a mandible fragment with c1 and p4-m1 and isolated M1 (Fig. 6D1, 2, Table 3); the age of the locality is MN13 (H ır and K€ okay, 2010). Eomellivora hungarica differs from E. piveteaui in its shorter mandible and reduced number of lower premolars, its larger size, and more robust dentition. The most remarkable feature of this species is the great robustness and length of its p4 . This tooth shows a well-marked backward inclination of the cuspids, with a clear imbrication with the mesial part of the m1. Also, the m1 has a strong central hypoconid with very marked lingual cristid. However, its M1 looks very similar to that of E. piveteaui, suggesting a robust form with some degree of affinity with E. piveteaui. Kretzoi (1942) also described E. hungarica altera from Cs akvar (Hungary), MN11 (H ır and K€ okay, 2010), based on three isolated teeth (P3, P4, and m1). Wolsan and Semenov (1996) synonymized it with E. wimani wimani. Despite substantial differences such as large upper dentition and a P3 with large lingual projection, there is not enough material to compare it with the other species of Eomellivora. We believe this material likely represents a distinct taxon, and yet because of the scarcity of the material and its lack of definitive diagnostic features, we assign them for now simply to Eomellivora sp. (Figs. 4,5,8, Table 3).
More recently, Morales and Pickford (2005) described the African E. tugenensis, from Ngorora Formation (Kenya), dated to about 12 Ma. This species is based on a fragmentary skull that contains a P4 with broken protocone and a complete M1. It is a primitive Eomellivora morphologically similar to E. wimani but much smaller (Werdelin and Peigne, 2010), being approximately intermediate in size between the extant G. gulo and M. capensis. The proportions of the P4 of E. tugenensis are similar to G. gulo, and thus very different than the other species of Eomellivora, but the proportions of the M1 are similar to the species of this genus (Figs. 4,5,7,8), suggesting that these African remains could correspond to an ancestral form of the genus, although more material is needed to address this hypothesis.

Phylogenetic Analysis
To better assess the relationships of E. piveteaui to other species of Eomellivora and within Mustelidae, we performed a cladistic analysis of nine taxa and 43 dental characters, being equally weighted and unordered (Figs. 7,9). The complete list of taxa, characters, and character-taxon matrix are available in Appendices 1 and 2. We restricted our analysis of the fossils to E. piveteaui from Batallones, E. wimani from Shangyingou, and E. ursogulo from Grebeniki-the only specimens with complete dental remains. Cladistic analysis was performed on this data set using PAUP*4.0b10 (Swofford, 2002). A bootstrap analysis was performed using 1000 replicates to test for resulting clade support. This analysis yielded a bootstrap tree of 98 steps, with a consistency index (CI) D 0.4592, a homoplasy index (HI) D 0.5408, and a retention index (RI) D 0.3977.
The cladogram shows that Eomellivora is the sister taxon of the extant Mellivora capensis (bootstrap 65%); thus, it is consistent to include it in the subfamily Mellivorinae (Fig. 9). This subfamily is characterized by a robust P3, with a high crown and marked buccal concavity, long relative to the P4; robust P4 with longer protocone; p4 relatively long and inclined backward; and m1 without metaconid, with a narrow talonid composed of a central hypoconid. Additionally, the monophyletic group Eomellivora is a well-supported clade (71% bootstrap support) ( Fig. 9) characterized by a relatively slender M1 with an enlarged stylar area, and a very swollen lingual cingulum; m1 trigonid occupying almost three-fourths of the total length of the tooth, with a high talonid composed of a buccolingually compressed hypoconid; and m2 without metaconid. E. wimani and E.ursogulo form a well-supported crown group (91% bootstrap support) (Fig. 9) characterized by a triangular P2 with a strong lingual cingulum and a buccal concavity on the wall; P3 with mesial accessory cuspid present and very marked mesial cingulum; lingual platform of M1 completely enclosing the protocone; p2 very robust; p3 with a buccolingual platform; and p4 with a high mesial accessory cuspid.

INSIGHTS INTO THE DIET OF EOMELLIVORA
The dentition of E. piveteaui has a mixture of features that can help us to interpret the feeding habits of this species. On the one hand, E. piveteaui possesses traits that can be interpreted as hypercarnivorous, based on features that emphasize the shearing function: (1) straight dental series; (2) long and buccolingually compressed m1 trigonid; and (3) loss of the m1 metaconid. All of these are features it shares with the extant Gulo gulo and Mellivora capensis and as well as with the extinct Ekorus ekakeran, Megalictis ferox, and the other species of Eomellivora; (4) the presence of a trenchant hypoconid centrally located on the m1 talonid, shared with all species of Eomellivora, that show a higher and more buccolingually compressed hypoconid than M. ferox and Ekorus ekakeran; and (5) m1 and m2 cuspids in line with the dental series, thereby increasing the length of the cutting edges. In contrast, Eomellivora piveteaui possesses other characteristics that could be interpreted as adaptations to durophagy and/or carcass processing: (1) p3-p4 blunt posterior accessory cuspids; (2) P4 protocone robust; (3) robust M1 lingual cingulum; and (4) distal widening of P3 and p3. Lungu (1978) considers E. piveteaui to be a smaller-sized Eomellivora with a more slender dentition than E. wimani, E. ursogulo, E. hungarica, and Eomellivora from Cs akv ar being a more primitive and less specialized species for crushing bones. Consequently, the complete sample of E. piveteaui from Batallones has allowed us to distinguish the existence of two dietary ecomorphotypes in the genus: (1) a group of smaller-size Eomellivora adapted to a more hypercarnivorous diet composed of E. piveteaui (Fig. 8), similar to that of the hypercarnivorous Ekorus ekakeran; and (2) a larger and robust Eomellivora composed of E.wimani, E. ursogulo, E. hungarica, and Eomellivora sp. from Cs akv ar characterized by having greater dimensions, and a higher and a greater number of blunt accessory cusps (Fig. 8), that may be adapted to a diet with a higher proportion of bone, similar to the extant durophage G. gulo (Larivi ere and Jennings, 2009) (although the higher number of accessory cusps would indicate greater hypercarnivory by some metrics; Hartstone-Rose, 2011; Hartstone-Rose and Stynder, 2013). The widening of some premolars, together with the more sectorial dentition in E. piveteaui, could be considered a hypercarnivorous specialization towards a more hypercarnivorous canid-like animal, capable of processing bones with its molars to maximize access to the available food.
Eomellivora piveteaui (Fig. 10) could have accessed a wide variety of prey items present in the fauna of Batallones. It likely occupied an ecological niche of an active predator of medium to large prey such as moschids, small-to medium-sized bovids, and young suids or cervids, while maintaining the ability to hunt small prey such as reptiles, birds, and lagomorphs. It may have occupied an intermediate ecological role between the opportunistic extant Mellivora capensis and the predominantly scavenging extant G. gulo. Moreover, E. piveteaui may have used its dentition to more fully process carcasses of its own prey, as happens with M. capensis, which is capable of consuming prey completely, including small bones (e.g., large lizards, snakes, or small mammals), but not so much as the efficient scavenger G. gulo (Ewer, 1998;Begg et al., 2003;Larivi ere and Jennings, 2009). Moreover, preliminary observations of the postcranial remains of E. piveteaui from Batallones show that the taxon had relatively long fore-and hind limbs, similar to extant canids that tend to have a high proportion of meat in their diet (e.g., Canis lupus). This could suggest possible cursorial adaptations in E. piveteaui. Future study of the postcranial remains could help determine whether this taxon was a pursuit predator and shed light on our understanding of its role in the Batallones ecosystem.

CONCLUSIONS
The craniomandibular features of the material from Batallones is assigned to the Eomellivora piveteaui, Ozansoy, 1965, providing a fuller picture of this species, supplementing the previously known material from Yassi€ oren and allowing the increased validation of the taxon at the specific level. The sample of E. piveteaui from Batallones permits the refinement of the diagnosis of the species, as well as expanding its age to MN10. Moreover, detailed comparisons with the rest of the species of Eomellivora show several morphological features that lead us to accept E. piveteaui, E. wimani, E. ursogulo, and E. hungarica. Eomellivora is the sister taxon of the extant Mellivora capensis, and E. piveteaui had a common ancestor with the crown group E. wimani-E. ursogulo. Eomellivora piveteaui was a large mustelid (Fig. 10) adapted to a more hypercarnivorous diet than the largest extant terrestrial mustelids (e.g., G. gulo and M. capensis), and the other species of Eomellivora and also shows some bone-crushing or carcass-processing adaptations. In addition, we suggest that E. piveteaui may have had an active predatory role, exploiting both small and relatively large prey.