This is the accepted version of the article: Alba, David M.; Robles Gimenez, Josep Maria; Valenciano Vaquero, Alberto; [et al.]. «A new species of Eomellivora from the latest Aragonian of Abocador de Can Mata (NE Iberian Peninsula)». Historical Biology, Published online July 2021. DOI 10.1080/08912963.2021.1943380 This version is available at https://ddd.uab.cat/record/247665 under the terms of the license A new species of Eomellivora from the latest Aragonian of Abocador de Can Mata (NE Iberian Peninsula) David M. Alba1,*, Josep M. Robles1, Alberto Valenciano2, Juan Abella1,3, Isaac Casanovas- Vilar1 1 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain 2 Departamento de Ciencias de la Tierra and Instituto Universitario de Investigación en Ciencias Ambientales de Aragón (IUCA), Área de Paleontología, Universidad de Zaragoza, C/ Pedro Cerbuna 12, 50009 Zaragoza, Spain 3 Instituto Nacional de Biodiversidad, Pje. Rumipamba N. 341 y Av. de los Shyris (Parque La Carolina) Quito, Ecuador *Corresponding author. E-mail address: david.alba@icp.cat ORCIDs: David M. Alba: 0000-0002-8886-5580 Josep M. Robles: 0000-0002-5410-3529 Alberto Valenciano: 0000-0003-1633-2248 Juan Abella: 0000-0002-3433-6093 Isaac Casanovas-Vilar: 0000-0001-7092-9622 1 ABSTRACT Eomellivora is a large-bodied mellivorine mustelid genus widely distributed throughout Eurasia and North America during the late Miocene (MN9-MN13). Here, we report the oldest Eurasian material of Eomellivora based on a palate and two mandibular fragments from ACM/PTA-A2, a pre-Vallesian (11.21 Ma; latest MN7+8) locality of Abocador de Can Mata (Vallès-Penedès Basin, NE Iberian Peninsula) that slightly predates the first appearance datum of Hippotherium by ~30 kyr. The described material differs from Hoplictis helbingi—another large mustelid recorded within the same basin in the roughly coeval site of Castell de Barberà (~11.2, earliest MN9)—and more closely resembles Eomellivora spp. Despite closer resemblances in both size and dental shape with the Vallesian (MN9–MN10) species Eomellivora piveteaui, the ACM material differs from multiple features that may be considered plesiomorphic. A new species, Eomellivora moralesi sp. nov., is thus erected based on the described material. A cladistic analysis confirms that the new species occupies a basal-most position within the Eomellivora clade, in agreement with its older age and more plesiomorphic morphology. KEYWORDS Carnivora, Mustelidae, Mellivorinae, Systematics, Late Miocene, Spain. 2 Introduction Eomellivora Zdansky, 1924 is a giant and hypercarnivorous mustelid genus recorded from the Miocene of Eurasia (Wolsan and Semenov, 1996; Valenciano et al. 2015, 2017) and North America (Baskin 1988). Wolsan and Semenov (1996) distinguished a single species of Eomellivora (the nominotypical one) with two distinct (Vallesian and Turolian) chronosubspecies. However, the recent description of additional material led Valenciano et al. (2015, 2017) to recognize as much as five species (see Table S1 for a list of localities). Older records of Eomellivora from the middle Miocene of Asia (e.g., Patnaik 2013) are either doubtful or correspond to species excluded from Eomellivora (Wolsan and Semenov 1996), whereas ?Eomellivora tugenensis Morales and Pickford, 2005 from Kenya (~12–11.5 Ma; Werdelin and Peigné 2010) most likely belongs to a different genus (Valenciano et al. 2017; Valenciano and Govender 2020), probably more closely related to the Mellivorini than to the Eomellivorini. Eomellivora sp. has also been reported from pre-Vallesian levels in Hammerschmiede 5 (Germany, ~11.62 Ma; Kirscher et al. 2016), but the material is unpublished. Here we report a new species of Eomellivora that further indicates that the genus was already present in Europe slightly before the Vallesian. Age and geological background The described material comes from the local stratigraphic sequence of Abocador de Can Mata (ACM; Alba et al. 2006, 2017) in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula; Figure 1a). Continuous paleontological surveillance during the construction of a dump since 2002 has yielded thousands of fossil vertebrate remains that can be accurately dated thanks to litho-, bio- and magnetostratigraphic correlation (Casanovas-Vilar et al. 2016a, 2016b; Alba et al. 2017). The published composite sequence 3 spans from 12.6 to 11.4 Ma (Alba et al. 2017), but additional fieldwork in 2019–2020 extended the sequence toward the Vallesian. The described material comes from locality ACM/PTA-A2 (Préstec de Terres de l’Abocador, subsector A, locality 2; Figure 1(b–c); Figure S1), which is situated in meter 268 of the ACM composite sequence and magnetostratigraphically correlated to the second portion of C5r.2n (i.e., postdating the short normal subchron C5r.2r-1), with an interpolated age 11.21 Ma and located only 5 m below the bottom C5r.1n (D.M.A. and J.M.R., unpublished data; see Figure S2). The beginning of the Vallesian in the Vallès-Penedès Basin, defined by the entry of hipparionin horses, is correlated to the base of C5r.1n, with an interpolated age of 11.18 Ma (Garcés et al. 1996; Casanovas-Vilar et al. 2016a, 2016b; Alba et al. 2019). Therefore, ACM/PTA-A2 can be securely correlated to latest Aragonian (MN7+8; Democricetodon crusafonti – Hippotherium Interval subzone of the Vallès-Penedès Basin; Casanovas-Vilar et al. 2016b), minimally predating the Vallesian by ~30 kyr. Material and methods The described material is housed at the Institut Català de Paleontologia Miquel Crusafont (ICP), Sabadell, Spain. It includes three dentognathic specimens recovered during April 2019 that might belong to a single individual, although the lack of contact and close spatial association makes it impossible to confirm this. Measurements were taken with a digital caliper to the nearest 0.1 mm. Maximum mesiodistal length (L) and buccolingual width (W) were measured and compared with data taken from the literature (Petter 1963; Wolsan and Semenov 1996 and references therein; Morales and Pickford 2005; Koufos 2012; Valenciano et al., 2015, 2017; Lavrov and Gimranov 2018) using bivariate plots. A robusticity index (in %) was also computed as RI=W/L × 100. 4 A cladistic analysis of Eomellivora spp. was performed based on a restricted subsample of the character-taxon matrix of Valenciano and Govender (2020; see character definition in Text S1), including three Eomellivora species, the mellivorin Howellictis valentini de Bonis et al., 2009, and the oligobunine Zodiolestes daimonelixensis Riggs, 1942 as outgroup. Two additional species of Eomellivora (based on Valenciano et al. 2017 and the Can Llobateres material described by Crusafont-Pairó 1972) and the new species described herein were also included in the analysis. The eomellivorin Ekorus ekakeran Werdelin, 2003 was excluded from the present analysis pending more detailed studies focused on the whole tribe. In turn, as in Valenciano and Govender (2020), the poorly known African eomellivorine ?E. tugenensis was excluded from the analysis because too few characters (15 out of 100) could be coded. Finally, the material from Gritsev assigned to Eomellivora wimani by Wolsan and Semenov (1996) and included in the morphometric comparisons as Eomellivora sp. was also excluded from the phylogenetic analysis pending its detailed description. The resulting matrix (Table 1) was analyzed using maximum parsimony with PAUP* v. 4.0a168 for Mac (Swofford 2003) with the ‘branch and bound’ option. All characters were treated as unordered and clade stability was assessed by means of bootstrap analysis (10,000 replicates) and Bremer indices. Systematic paleontology Order Carnivora Bowdich, 1821 Suborder Caniformia Kretzoi, 1943 Family Mustelidae Fischer, 1817 Subfamily Mellivorinae Gray, 1865 Tribe Eomellivorini Zdansky, 1924 5 Genus Eomellivora Zdansky, 1924 Emended diagnosis Modified after Valenciano et al. (2017). Mellivorine mustelid of large size; P1 present; P3 with the distal area thickened (only incipiently in the most plesiomorphic species); P4 with a subconical protocone, and with paracone-protocone and paracone-parastyle crests; P4 protocone located in line with the parastyle; P4 parastyle poorly-developed but thickened; buccal wall of P4 with a concavity in the base of the crown between the paracone and the metastyle, exhibiting a variable degree of development; stylar area of M1 enlarged; M1 with a non-reduced metacone in the more plesiomorphic species and a reduced one in the more derived ones; M1 with an arched ridge-shaped or conical protocone continuing into the mesial protocone crest, and a talon relatively equally expanded mesially and distally; premolar teeth crowned; p1 present; p2 turned buccolingually from the tooth row; p3 with a distal accessory cuspid and with the distal area thickened; p4 enlarged with a distal accessory cuspid and with a backward inclination of the main cuspid towards the m1; m1 enlarged with a distinct metaconid in the most plesiomorphic species and vestigial to absent in the remaining ones, in which it is replaced by a distinct crest; m1 talonid with single but strong, high and centrally positioned hypoconid; m2 elongated mesiodistally with a low crown surrounded by a cingulum and a central protoconid. Included species Eomellivora wimani Zdansky, 1924 (type species); Eomellivora fricki (Pia, 1939); Eomellivora hungarica Kretzoi, 1942; Eomellivora ursogulo (Orlov, 1948); Eomellivora piveteaui Ozansoy, 1965; Eomellivora moralesi sp. nov. 6 Eomellivora moralesi sp. nov. Holotype IPS122262, palate with incisor, canine and left P1 alveoli, as well as left P2–M1 and right P1–M1 crowns (Figure 2(a–e, l–s)). Paratypes IPS122212, right mandibular fragment preserving the symphysis, c1–p1 alveoli, the complete p2 crown, and partial p3 (very damaged) and p4 (only mesial) crowns (Figure 2(f– h)); IPS122214, right mandibular fragment with m2 (Figure 2(i–j)). Type locality ACM/PTA-A2 (els Hostalets de Pierola, Catalonia, Spain). Age and distribution Only known from the type locality (11.21 Ma, latest Aragonian, MN7+8). Etymology Species trivial name dedicated to Prof. Jorge Morales, in recognition to his extraordinary contribution to vertebrate paleontology. Diagnosis 7 Eomellivora species of moderate size (similar to E. piveteaui). P2 without distal accessory cusp and with a subrectangular occlusal profile that lacks a labial concavity. P3 slender, with a straight labial wall, a poorly developed basal cingulum, and without mesial and distal accessory cusps. P4 without a marked concavity on the labial wall or a marked basal cingulum. M1 slender, with a well-developed parastylar area and lingual platform (longer than the labial wall) that lacks a concavity at mid-length, a non-reduced metacone as well as a mesiolingually located protocone that bears a ridge-like extension and is not completely enclosed by the lingual platform. p2 moderately robust and without distal accessory cuspid. p3 moderately expanded distally. m2 with metaconid. The new species differs from other Eomellivora spp. in the slenderer P3 without a distal accessory cusp, a marked labial concavity or a strong basal cingulum (tooth locus unknown in E. fricki and E. hungarica); the weaker P4 basal cingulum (tooth locus unknown in E. hungarica); and the retention of m2 metaconid (unknown in E. fricki and E. hungarica). It further differs from E. piveteaui in the larger parastylar area of the M1, and the subquadrangular lingual platform of the M1, and from E. fricki in the smaller size, the lower P4 protocone, and the less robust M1 with a less developed lingual platform. The new species more clearly differs from E. wimani, E. ursogulo, and E. hungarica in the more mesially located protocone and the non-reduced metacone (relative to the paracone) in the M1, and the lack of p4 mesial accessory cuspid. It further differs from E. wimani and E. ursogulo in other features unknown for E. hungarica: the subrectangular (instead of triangular) occlusal profile of the P2 with a straight (non-concave) labial wall, the lack of P3 mesial accessory cusp, the lack of a marked labial concavity on the P4, and the less robust p2; additionally from E. wimani, in the more ridge-like M1 protocone; and additionally from E. ursogulo, in the lack of distal accessory cusp in the P2, of a mid-length concavity on the 8 M1 lingual platform, and of a distal accessory cuspid in the p2, as well as in the less distally expanded p3. Description Craniodental measurements for the holotype and paratypes are reported in Tables 2–3 (see also Figure 3 for bivariate plots of tooth dimensions). The cranial fragment IPS122262 (Figure 2(a–e)) preserves the palate and the anteriormost portion of the zygomatic arches, which are crushed and held together by matrix. The specimen displays an overall massive aspect and preserves relatively well the original shape of the dental arcade. The infraorbital foramen is large (Table 3) and located at the level of the upper carnassial parastyle. The external border of the right C1 is broken away and both the right premaxilla and palatine process of the maxilla at the I3–P2 level are somewhat displaced from their anatomical position, but it can be ascertained that the right foramen incisivum would have been located at the mesial level of the C1 alveolus. No incisors are preserved, but the I1–I2 alveoli are transversely aligned and overlap with the anteriormost portion of the I3 alveolus. Based on alveolus size, the I3 crown would have been caniniform and much larger than the other incisors, but about half the size of the C1, which appears quite massive and only slightly labiolingually compressed in basal dimensions (estimated RI = 94%). There is a small diastema between the I3 and the C1. Between the distolingual corner of the C1 alveolus and the P2 there is a unicuspid, uniradiculate and tiny P1, whose crown is only preserved on the right side. The P2 crown (Figure 2(l–m)) is biradiculate and displays a very slender (RI=61–62%), elliptical to subrectangular occlusal outline (much wider than long), with a straight labial and convex lingual sides. The P2 is rotated labially, its main cusp is mesially located, and it lacks a 9 distinct distal accessory cusp. The P3 (Figure 3(n–o)) is much larger than the P2. It is biradiculate and displays a subrectangular but slenderer contour (RI=55–56%) than the P2. The P3 lingual margin is bulging, whereas the labial wall is straight. This tooth has marked mesial and distal cingula, which nevertheless do not continue along the lingual or labial walls. There are no distinct mesial or distal accessory cusps, and indeed the thick and blunt crista that extends distalward from the main cusp apex fades away before reaching the distal margin. The P4 (Figure 2(p–q)) is longer than the P3 (P4 L / P3 L = 147–149%) and also somewhat stouter than the preceding premolars (RI = 65–66%). The occlusal morphology is best preserved in the right P4, while the left one is distally damaged. There is some dentine exposure at the apices of the protocone and the paracone, as well as along the metastyle, which extends along about two-thirds of crown length. The crown displays a subtriangular occlusal profile with almost parallel (non-distally tapering) labial and lingual walls. The protocone is low and peripherally located on the protruding mesiolingual corner of the crown, being surrounded by a distinct cingulum. The parastyle is low and transversely aligned with the protocone. The crown displays a very slight mesial concavity between parastyle and protocone, and an almost straight buccal crown wall (with no distinct concavity distally from the paracone). The paracone is the highest and most extensive cusp; together with the metastyle it constitutes a moderately elongate, slightly labially concave blade. A distinct mesiolabial crista extends from close to the paracone apex to the parastyle, whereas a fainter mesiolingual crista links the paracone with the protocone. There is no well-developed cingulum surrounding the whole crown. The M1 (Figure 2(r–s)) has a vaguely figure-eight shape, being considerably broader than long (RI=173–174%) but slightly narrower than the length of the upper carnassial (M1 W / 10 P4 L = 89–90%). The crown is markedly constricted at about mid-width, with an expanded lingual platform that displays a suboval (convex) profile and is much longer than the buccal half of the crown. The tooth is surrounded by a marked cingulum, well developed on the lingual side and on the labial side, where it constitutes a distinctly protruding (almost cuspule-like) stylar area. The paracone is bulbous and more extensive than the metacone, which is partially worn. The protocone is pyramidal but displays a ridge-like mesiolabial extension. It is centrally located on the labial moiety of the crown and is not completely enclosed within the lingual platform. There is no conspicuous crest linking the paracone with the protocone. Based on IPS122212 (Figure 2(f–h)) and IPS122214 (Figure 2(i–k)), the morphology of the mandible can be only partially ascertained. The former specimen preserves most of the symphyseal suture (from the mesialmost level of the c1 alveolus) and the corpus up to mid- p4 level, whereas the latter preserves the posteriormost portion of the corpus (including the distal portion of the m1 alveolus) and the anteriormost portion of the ramus. The symphysis displays a gently convex anterior profile and inferiorly extends up to the p3 level, where the corpus is somewhat deeper than below the p4. Overall, the corpus appears dorsoventrally deep and labiolingually thick, being stouter at the c1 level than posteriorly (Table 3). There are two mental foramina slightly above corpus mid-height, one below the p2 and the other at the distal p3 level. There is a deep masseteric fossa that reaches its maximum depth at the distal m2 level but extends rostrally beyond the m2. Based on alveolus size, the c1 would have been quite massive, displaying a slightly labiolingually compressed subelliptical basal contour (estimated RI=77%). There is a tiny p1 alveolus between the mesiolingual aspect of the p2 and the distal aspect of the c1 alveolus. The p2 (Figure 2(t)) is biradiculate and much longer than wide (RI=58%), with its main axis 11 labially rotated. There is no distinct distal accessory cuspid and there are no cingula other than the thickened distal crown margin. Based on its alveolus, the p3 would have been somewhat rotated labially, considerably longer than the p2 (12.4 vs. 8.7 mm, respectively), and slightly slenderer (estimated RI=52%), being wider distally than mesially. Although only the distalmost portion of the p3 is preserved, a distal accessory cuspid seems to have been present on the distolabial aspect of the main cuspid. Only the mesial portions of the p4 crown (including the apex of the main cuspid) and alveolus are preserved, making it impossible to ascertain the size and proportions of this tooth. There is no mesial accessory cuspid and the preserved portion of the labial wall appears straight. The size and proportions of the m1 cannot be adequately ascertained either. The m2 (Figure 2(u)) is obliquely inserted (relative to the occlusal plane of the remaining postcanine teeth). It displays a trapezoidal occlusal contour, being slightly longer than broad (RI=84%), particularly on the labial side, and possessing a markedly oblique distal margin. The protoconid is rounded and centrally located, but the mesial aspect of the m2 is worn away, making it impossible to ascertain the development of the paraconid. In contrast, a low but extensive and distally protruding hypoconulid is present on the distolabial corner of the crown. A distinct (albeit smaller) metaconid is also present on the lingual aspect of the protocone base. Phylogenetic results The phylogenetic analysis based on 24 parsimony-informative characters yielded 3 most parsimonious cladograms of 34 steps (Figure 4). The analysis fails to resolve the trichotomy between E. wimani, E. ursogulo, and E. hungarica but recovers E. moralesi sp. nov. as the 12 most basal species of the genus, followed by the successive branching of E. fricki and E. piveteaui. Discussion and conclusions Eomellivora moralesi sp. nov. from ACM generally fits Valenciano et al.’s (2017) emended diagnosis of Eomellivora (see also Figure 5), except for the presence of one or two distal accessory cusps in the P3. The ACM material further differs from other Eomellivora species in the retention of a metaconid in the m2. Accordingly, an emended diagnosis of the genus (modified after Valenciano et al. 2017) has been provided. Both features are probably plesiomorphic, in agreement with the somewhat older age of the new species and its greater similarities with the Vallesian species of Eomellivora (e.g., subrectangular P2 crown without a labial concavity; P3 without mesial accessory cusp; lack of a marked concavity on the labial wall of the P4; non-reduced M1 metacone; moderately robust p2; lack of p4 mesial accessory cuspid). The retention of plesiomorphic features in E. moralesi sp. nov. agrees with our cladistic results indicating a basalmost position within the genus. ?Eomellivora tugenensis from Kenya (Morales and Pickford 2005), potentially ancestral to later Eomellivora (Valenciano et al. 2015, 2017), was excluded from the analysis because of excessive missing data. The small size, relatively large orbit size and lack of P1 suggest that ?E. tugenensis might be more closely related to mellivorins than to eomellivorins. In any case, the ACM material clearly differs from it by the larger size, the relatively smaller orbit, the more transversely aligned incisors, the less labiolingually compressed C1, the presence of a small diastema between the I3 and the C1, the presence of P1, the rotated orientation of the P2, the more robust P4, and the more distinct labial stylar area in the M1. 13 An alternative assignment of the ACM material to Hoplictis Ginsburg, 1961 can also be discounted based on the larger size of the former, the stouter P4 with a better developed protocone, and the non-reduced metacone and larger lingual platform of the M1 (Valenciano et al. 2019). Only Hoplictis helbingi (Viret, 1951), known by an m1 (lectotype; Depéret 1892: Pl. I fig. 3; Viret, 1951: Pl. II fig. 12a-b) and a P4 (paralectotype; Viret 1951: Pl. II fig. 13a–b) from La Grive-Saint-Alban (MN7+8), France, approaches the size of the ACM material. The lectotype of H. helbingi is virtually identical to that of a mandibular fragment (IPS33185) from Castell de Barberà (~11.2 Ma; Alba et al. 2019) assigned to the same species (Crusafont-Pairó 1972: Pl. 1 fig. 1; Valenciano et al. 2019: fig. 4l–n). Compared to the latter, the ACM material displays a deeper and stouter mandibular corpus, with the m2 alveolus being socketed obliquely (as it is typical of Eomellivora) instead of being on the same alveolar plane as the m1. The P4 from ACM further differs from the paralectotype of H. helbingi by being slightly larger and more robust, displaying parallel (instead of distally tapering) labial and lingual margins, and possessing a relatively larger and more mesially protruding protocone. Eomellivora had not been reported as such from the Vallès-Penedès Basin until Valenciano et al. (2019) synonymized Hoplictis petteri (Crusafont-Pairó, 1972) with E. fricki based on the P4 holotype of the former (Fig. 5k) from Can Llobateres 1 (9.76 Ma; Casanovas-Vilar et al. 2016b), originally described by Petter (1963). The scarce material from Los Valles de Fuentidueña (MN9; Crusafont-Pairó and Ginsburg 1973; Ginsburg et al. 1981), previously assigned to E. piveteaui (Wolsan and Semenov 1996; Valenciano et al. 2015, 2017), might also be referrable to E. fricki based on the great development of the lingual platform and the distal expansion of the metacone in the M1 (Fig. 5r). Coupled with the material from Austria (Pia 1939; Zapfe 1948; Valenciano et al. 2017), this suggests that E. 14 fricki was widely distributed across Europe during the early Vallesian. However, an assignment of the ACM material to E. fricki can be ruled out based on its smaller size (Fig. 3) and other occlusal differences (Fig. 5). The ACM material is indeed more similar in size and occlusal morphology to E. piveteaui (Figs. 3 and 5), which is also known from the Spanish Vallesian (Batallones-3 and 10, MN10; Valenciano et al., 2015). Nevertheless, the ACM material from ACM differs from both E. piveteaui and E. fricki in multiple occlusal features present also in other Eomellivora (Figure 5), indicating that it belongs to a new, more plesiomorphic species. The somewhat older age of the new species further indicates that Eomellivora dispersed into (or locally evolved in) Europe sometime before the dispersal of hipparionin horses. Acknowledgments We thank CESPA Gestión de Residuos S.A.U. for defraying fieldwork; the ICP Preparation & Conservation Area for preparing the specimens; Víctor Vinuesa and Itzíar Llopart for codirecting fieldwork; the Centre d’Interpretació i Restauració Paleontològica (els Hostalets de Pierola) and the Servei d’Arqueologia i Paleontologia of the Generalitat de Catalunya for their collaboration; our colleagues M. Sotnikova (PIN, Moscow), the late S. Peigné, and P. Loubry (MNHN, Paris) for kindly providing us photographs of the holotype of E. ursogulo and the lectotype of E. piveteaui; the reviewers Michael Morlo and Louis de Bonis for helpful comments that helped to improve a previous version of this paper; and the editor (Gareth Dyke) and the guest editors for inviting us to contribute to this special issue in honor of Jorge Morales. We wholeheartedly dedicate this paper to him: many thanks, Jorge, for being an endless source of inspiration! 15 Disclosure statement No potential conflict of authors was reported by the authors. Funding This work was supported by the Agencia Estatal de Investigación (CGL2016-76431-P, CGL2017-82654-P and PGC2018-094122-B-100, AEI/FEDER, EU); the Ministerio de Economía y Competitividad (RYC-2013-12470 to I.C.V.); the Generalitat de Catalunya/CERCA Programme; AGAUR (2017 SGR 116 and Beatriu de Pinós 2017 BP 00223 to J.A.); the Gobierno de Aragón (Group ref. E33_20R to A.V.); and the “Juan de la Cierva Formación” program from the Spanish Ministry of Science, Innovation, and Universities (FJC2018- 036669-I to A.V.). References Alba DM, Moyà-Solà S, Casanovas-Vilar I, Galindo J, Robles JM, Rotgers C, Furió M, Angelone C, Köhler M, Garcés M, et al. 2006. Los vertebrados fósiles del Abocador de Can Mata (els Hostalets de Pierola, l’Anoia, Cataluña), una sucesión de localidades del Aragoniense superior (MN6 y MN7+8) de la cuenca del Vallès-Penedès. Campañas 2002-2003, 2004 y 2005. Estud Geol. 62:295–312. Alba DM, Casanovas-Vilar I, Garcés M, Robles JM. 2017. Ten years in the dump: An updated review of the Miocene primate-bearing localities from Abocador de Can Mata (NE Iberian Peninsula). J Hum Evol. 102:12–20. Alba DM, Garcés M, Casanovas-Vilar I, Robles JM, Pina M, Moyà-Solà S, Almécija S. 2019. Bio- and magnetostratigraphic correlation of the Miocene primate-bearing site of Castell de Barberà to the earliest Vallesian. J Hum Evol. 132:32–46. 16 Baskin JA. 1998. Mustelidae. In: Janis CM, Scott KM, Jacobs LL, editors. Evolution of Tertiary mammals of North America. Volume 1: Terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press; p. 152–173. Bowdich TE. 1821. An analysis of the natural classifications of Mammalia, for the use of students and travellers. Paris: J. Smith. Casanovas-Vilar, I., Madern A., Alba DM, Cabrera L, García-Paredes I, Van den Hoek Ostende LW, DeMiguel D, Robles JM, Furió M, Van Dam J, et al. 2016a. The Miocene mammal record of the Vallès-Penedès Basin (Catalonia). C R Palevol. 15:791–812. Casanovas-Vilar I, Garcés M, Van Dam J, García-Paredes I, Robles JM, Alba DM. 2016b. An updated biostratigraphy for the late Aragonian and the Vallesian of the Vallès-Penedès Basin (Catalonia). Geol Acta. 14:195–217. Crusafont-Pairó, M. 1972. Les Ischyrictis de la transition Vindobonien-Vallésien. Palaeovertebrata. 5:253–260. Crusafont-Pairó M, Ginsburg L. 1973. Les carnassiers fossiles de Los Vallès de Fuentiduena (Ségovie, Espagne). Bull Mus Natl Hist Nat. 131:29–45. de Bonis L, Peigné S, Guy F, Likius A, Makaye HT, Vignaud P, Brunet M. 2009. A new mellivorine (Carnivora, Mustelidae) from the Late Miocene of Toros Menalla, Chad. N Jb Geol Paläontol. 252:33–54. Depéret C. 1892. La faune de Mammifères Miocènes de La Grive-Saint-Alban (Isère) et de quelques autres localités du bassin du Rhone. Arch Mus Hist Nat Lyon. 5:1–95. Fischer G. 1817. Adversaria zoologica. Mem Soc Imp Natural Moscou. 5:357–472. Garcés M, Agustí J, Cabrera L, Parés JM. 1996. Magnetostratigraphy of the Vallesian (late Miocene) in the Vallès-Penedès Basin (northeast Spain). Earth Planet Sci Lett. 142:381– 396. 17 Ginsburg L. 1961. La faune des Carnivores Miocènes de Sansan (Gers). Mém Mus Natl Hist Nat. 9:1–190. Ginsburg L, Morales J, Soria D. 1981. Nuevos datos sobre los carnívoros de los Valles de Fuentidueña (Segovia). Estud Geol. 37:383–415. Gray JE. 1865. Revision of the genera and species of Mustelidae contained in the British Museum. Proc Zool Soc Lond. 1865:100–154. Kirscher U, Prieto J, Bachtadse V, Aziz HA, Doppler G, Hagmaier M, Böhme M. 2016. A biochronologic tie-point for the base of the Tortonian stage in European terrestrial settings: Magnetostratigraphy of the topmost Upper Freshwater Molasse sediments of the North Alpine Foreland Basin in Bavaria (Germany). Newslett. Stratigr. 49:445–467. Koufos GD. 2012. New material of Carnivora (Mammalia) from the Late Miocene of Axios Valley, Macedonia, Greece. C R Palevol. 11:49–64. Kretzoi M. 1942. Eomellivora von Polgárdi und Csákvár. Föld Közl. 72:318–323. Kretzoi M. 1943. Kochictis centennii n. g. n. sp., ein altertümlicher Creodonte aus dem Oberoligozän Siebenbürgens. Föld Közl. 73:190–195. Lavrov AV, Gimranov DO. 2018. First finding of a representative of giant mustelids of the genus Eomellivora (Carnivora, Mustelidae) in Russia (Tuva, Upper Miocene). Doklady Biol Sci. 480:82–84. Morales J, Pickford M. 2005. Carnivores from the Middle Miocene Ngorora Formation (13- 12 Ma), Kenya. Estud Geol. 61:271–284. Orlov YU. 1948. Perunium ursogulo Orlov, a new gigantic extinct mustelid (a contribution to the morphology of the skull and brain and to the phylogeny of Mustelidae). Acta Zool. 29:63–105. 18 Ozansoy F. 1965. Étude des gisements continentaux et des mammifères du Cénozoïque de Turquie. Mém Soc Géol Fr. 44:1–92. Patnaik, R. 2013. Indian Neogene Siwalik mammalian biostratigraphy. An overview. In: Wang X, Flynn LJ, Fortelius M, editors. Fossil mammals of Asia: Neogene biostratigraphy and chronology. New York: Columbia University Press; p. 423–444. Petter G. 1963. Contribution a l'étude des Mustélidés des bassins néogènes du Vallès- Pénédès et de Calatayud-Teruel. Mem Soc Géol Fr. 97:5–44. Pia J. 1939. Ein riesiger Honingsdachs (Mellivorine) aus dem Unterpliozän von Wien. Ann Naturhist Mus Wien. 50:537–583. Riggs ES. 1942. Preliminary description of two Lower Miocene carnivores. Field Mus Nat Hist Geol Ser. 8:59–62. Swofford DL. 2003. PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Version 4. Sunderland: Sinauer Associates. Valenciano A, Abella J, Sanisidro O, Harstone-Rose A, Álvarez-Sierra MÁ, Morales J. 2015. 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). J Vertebr Paleontol. 35:e934570. Valenciano A, Abella J, Göhlich UB, Ángeles Álvarez-Sierra M, Morales J. 2017. Re-evaluation of the very large Eomellivora fricki (Pia, 1939) (Carnivora, Mustelidae, Mellivorinae) from the late miocene of Austria. Palaeontol Electron. 20:17A. Valenciano A, Jiangzuo Q, Wang S, Li C, Zhang X, Ye J. 2019. First record of Hoplictis (Carnivora, Mustelidae) in East Asia from the Miocene of the Ulungur River Area, Xianjiang, Northwest China. Acta Geol Sin. 93:251–264. 19 Valenciano A, Govender R. 2020. New fossils of Mellivora benfieldi (Mammalia, Carnivora, Mustelidae) from Langebaanweg, 'E' Quarry (South Africa, Early Pliocene): Re-evaluation of the African Neogene mellivorines. J Vertebr Paleontol. 40:e1817754. Viret J. 1951. Catalogue critique de la faune des Mammifères Miocènes de la Grive Saint- Alban (Isère). Première Partie. Chiroptères, Carnivores, Édentés Pholidotes. Nouv Arch Mus Hist Nat Lyon. 3:1–102. Werdelin L. 2003. Mio-Pliocene Carnivora from Lothagam, Kenya. In: Leakey MG, Harris JM, editors. Lothagam: The dawn of humanity in Eastern Africa. New York: Columbia University Press; p. 261–328. Werdelin L, Peigné S. 2010. Carnivora. In: Werdelin L, Sanders WJ, editors. Cenozoic mammals of Africa. Berkeley: University of California Press; p. 603–657. Woslan M, Semenov YA. 1996. A revision of the late Miocene mustelid carnivoran Eomellivora. Acta Zool Cracov. 39:593–604. Zapfe H. 1948. Neue Funde von Raubtieren aus dem Unterpliozän des Wiener Beckens. Sitz- Ber Öst Akad Wiss math-naturwiss Kl. 157:243–262. Zdansky O. 1924. Jungtertiäre carnivoren Chinas. Palaeontol Sin C. 2:1–149. Figure captions Figure 1. a, Simplified geological map of the Vallès-Penedès Basin (modified from Casanovas-Vilar et al. 2016a: fig. 1); the location of Abocador de Can Mata is denoted by a black dot, whereas the top left inset denotes the location of the Vallès-Penedès Basin within the Iberian Peninsula. b–c, Aerial photographs showing the location of locality ACM/PTA-A2 (geographic coordinates ETRS89: 1.799865E, 41.528243N) within the area affected by the 20 landfill construction (in 2017 and 2020) and in relation to farmhouses and the locality ACM/BCV1 (b), as well as relative to subsector ACM/PTA-A and the farmhouse of Can Mata de la Garriga (c). Base orthophotos were taken in 2019 (b) and 2021 (c); © Institut Cartogràfic i Geològic de Catalunya, downloaded from VISSIR v. 3.26 and reproduced with permission by means of a Creative Commons license CC BY 4.0 (ICGC, 2021). Abbreviations: ACM = Abocador de Can Mata; BCV = Barranc de Can Vila; PTA = Préstec de Terres de l’Abocador. Figure 2. Eomellivora moralesi sp. nov. from ACM/PTA-A2: a–e, IPS122262 (holotype), palate in inferior (a), rostral (b), caudal (c), left lateral (d), and right lateral (e) views; f–h, IPS122212 (paratype), right mandibular fragment with symphysis and premolars, in occlusal (f), labial (g), and lingual (h) views; i–k, IPS122214 (paratype), right mandibular fragment with m2, in occlusal (i), labial (j), and lingual (k) views; l–u, complete tooth crowns of the aforementioned specimens in greater occlusal detail: l, rigth P2; m, left P2; n, right P3; o, left P3; p, right P4; q, left P4; r, right M1; s, left M1; t, right p2; u, right m2. Images are oriented with mesial (a, f, I, l–u) or dorsal (b–e, g–h, j–k) on top. Figure 3. Bivariate plots of buccolingual width (W, in mm) vs. mesiodistal length (L, in mm) to compare dental size and proportions in Eomellivora moralesi sp. nov. with those of other species of Eomellivora. Comparative data were taken from the literature (see Materials and methods for data sources). 21 Figure 4. Strict consensus of 3 most parsimonious cladograms of 34 steps. Consistency index = 0.735; retention index = 0.743; rescaled consistency index = 0.546. Bremer and bootstrap (when >50%) support for ingroup clades are depicted above and below nodes, respectively. Figure 5. Occlusal views of the P3 (a–d), P4 (e–k) and M1 (l–r) of Eomellivora species compared (all teeth depicted as from the right side): a, e, l) Eomellivora moralesi sp. nov. from ACM/PTA-A2 (IPS122262, holotype; P3 reversed); b, f, m) Eomellivora piveteaui from Batallones-3 (Bat-3´13.185; P3 reversed); c, g, n) Eomellivora ursogulo from Grebeniki (PIN- No.268, holotype); d, h, o) Eomellivora wimani from Shangyingou (PMU-M3692, lectotype); i, p) E. piveteaui from Yassiören (MNHN-TRQ-1005, holotype); j, q) Eomellivora fricki from Wien XII-Altmannsdorf (NHMW 2016/0065/0001, holotype; reversed); k) E. fricki from Can Llobateres 1 (IPS2015); r) E. fricki from Los Valles de Fuentidueña (IPS2057). All images are depicted with mesial on top. 22 A 6 D 18 W 5 P1 C 12 17 P4 P3 4 11 16 3 10 15 W 2 3 4 5 6 W L 9 14 B 7 8 13 P2 6 7 12 12 13 14 15 16 17 18 20 21 22 23 24 25 W E 5 L L 4 23 6 7 8 9 10 11 M1 G L 22 F 21 8 8 m2 p2 20 7 W 7 19 6 W 6 W 18 5 5 17 4 4 9 10 11 12 13 14 15 6 7 8 9 10 6 7 8 9 10 11 12 L L L E. moralesi sp. nov. E. fricki E. ursogulo Eomellivora sp. (Gritsev) E. piveteaui E. wimani E. hungarica Eomellivora sp. (Csákvár) Zodiolestes daimonelixensis Howellictis valentini Eomellivora moralesi 3 Eomellivora fricki 79 1 Eomellivora piveteaui 64 1 Eomellivora hungarica 55 2 Eomellivora wimani 84 Eomellivora ursogulo Table 1. Character-taxon matrix used in the phylogenetic analysis, following Valenciano & Govender (2020). See also Supplementary Text S1 for the description of character states. Only parsimony-informative characters for this sample of taxa (bolded) were used in the analysis. 0000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000001 Character # 0000000001111111111222222222233333333334444444444555555555566666666667777777777888888888899999999990 Taxon 1234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890 Eomellivora_moralesi ?110???1??0100000100?101010100111010001000100011???1?0????????????????????0?0??????????????????????? Eomellivora_fricki ??1?11?1???????0???????11?01011110?00120?01?0???????1000101?10?10??11?201?0????????????????????????? Eomellivora_hungarica ???????1???????????????????????111?10110????0???????1100101010111111122010?????????????????????????? Eomellivora_piveteaui 11101101??010001001111010101011110100010001000110001100010101011111111101000101000000111110110002101 Eomellivora_wimani ??10???1010101111011110101011111111111100010002100021100101010111111111010001??????????????????????? Eomellivora_ursogulo 1?1????1110111111011110101011111111101110011002110021100101010011111111010001??????????????????????? Howellictis_valentini 1?0?1101??100100000???11001000120111011001100021000110001010000111011110100110?????????????????11??? Zodiolestes_daimonelixensis 1100110001000000000001010011002110000100000100100001100000000000110000000000010001101000020000001010 Table 2. Dental measurements of Eomellivora moralesi sp. nov. from ACM/PTA-A2 based on the holotype (upper teeth) and paratypes (lower teeth). Measurements within parentheses are estimated due to minor damage; those within brackets are estimated based on alveolus size. Left Right Tooth L (mm) W (mm) W/L (%) L (mm) W (mm) W/L (%) I3 [6.0] [7.4] [123.3] — — — C1 [12.8] [12.0] [93.8] — — — P1 — — — 4.1 3.2 78.0 P2 8.0 4.9 61.3 8.1 5.0 61.7 P3 14.3 7.9 55.2 14.2 7.9 55.6 P4 (21.0) 13.8 (65.7) 21.1 13.8 65.4 M1 10.8 18.8 174.1 10.8 18.7 173.1 c1 [13.0] [10.0] [76.9] — — — p1 [3.0] [2.3] [76.7] — — — p2 8.7 5.0 57.5 — — — p3 [12.4] 6.4 [51.6] — — — m2 7.6 6.4 84.2 — — — Table 3. Cranial and mandibular measurements of Eomellivora moralesi sp. nov. from ACM/PTA-A2 based on the holotype (cranium) and paratypes (mandible). Variable Measurement (mm) Length of the preserved cranium 99 Width of the preserved cranium 122 Right infraorbital foramen width 9.8 Right infraorbital foramen height 7.5 Length of I3–C1 diastema 4.5 Mandibular corpus depth below p3 28.7 Mandibular corpus depth below p4 26.3 Mandibular corpus maximum width at c1 18.0 Mandibular corpus maximum width at p3 14.5 Mandibular corpus maximum width at p4 13.5 Supplementary material of “A new species of Eomellivora from the latest Aragonian of Abocador de Can Mata (NE Iberian Peninsula)” David M. Alba1,*, Josep M. Robles1, Alberto Valenciano2, Juan Abella1,3, Isaac Casanovas-Vilar1 1 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain 2 Departamento de Ciencias de la Tierra and Instituto Universitario de Investigación en Ciencias Ambientales de Aragón (IUCA), Área de Paleontología, Universidad de Zaragoza, C/ Pedro Cerbuna 12, 50009 Zaragoza, Spain 3 Instituto Nacional de Biodiversidad, Pje. Rumipamba N. 341 y Av. de los Shyris (Parque La Carolina) Quito, Ecuador *Corresponding author. E-mail address: david.alba@icp.cat 1 Figure S1. General (a) and detailed (b) views of locality ACM/PTA-A2 during excavation in May and June 2019, respectively. © Institut Català de Paleontologia Miquel Crusafont. 2 Figure S2. Stratigraphic section of Abocador de Can Mata, sector Préstec de Terres de l’Abocador, subsector A (ACM/PTA-A). Uppercase letters to the left correspond to lithostratigraphic correlation levels, whereas the stratigraphic situation of locality ACM/PTA-A2 (bottom of level N) is indicated with an arrow. Lithology abbreviations: C = claystones; S = siltstones; FS = fine sandstones; CS = coarse sandstones; CG = conglomerates. Note that the previously available composite lithostratigraphic correlation panel for ACM was based on 42 correlation levels (Alba et al. 2017: SOM Fig. S1 and Table S1), of which the uppermost one was level G. The whole stratigraphic section of ACM/PTA-A comprises correlation levels H to N, and is thus located above the ACM composite sequence reported by Alba et al. (2017). The distance between correlation levels G and H is ca. 10 m. Based on currently available data (D.M.A. and J.M.R.’s unpublished data), ACM/PTA-A2 is located in meter 268 of the updated composite sequence, being correlated to C5r.2n with an interpolated age of 11.21 Ma. 3 Supplementary Text S1. Characters and character states used in the phylogenetic analysis, after Valenciano & Govender (2020). (1) Posterior lacerate and jugular foramina: “fused” into a single foramen (0); tendency to separate openings with the jugular foramen distolateral to the posterior lacerate foramen (1). (2) Shape of upper incisor row: parabolic (0); straight (1). (3) Position of the infraorbital foramen: above P3 (0); above P4 parastyle (1). (4) Incisive foramen: located at the level of C (0); located at the level of the diastema I3-C (1). (5) Mastoid process: reduced, located in dorsal view in line with the middle point of the orbit (0); enlarged, located laterally exceeding to the orbit (1). (6) Relative position of the mastoid and paroccipìtal processes: relatively close (0); mastoid process located mesially to the paroccipital process (1). (7) Paroccipital process: not reduced (0); reduced (1). (8) Height of the mandibular corpus: low and thin mandibular corpus (0); high and thick mandibular corpus (1). (9) Orientation of the I3 cusp: spreader out laterally (0); in line with the cusps of I1-2 (1). (10) Orientation of Canine: spreader out laterally, with an arrangement of the tip none parallel (0); ventrally directed, with a parallel arrangement of the tip (1). (11) P1. Present (0); absent (1). (12) P2. Mesiodistal axis of P2: in line with the tooth row (0); rotated buccolingually (1). (13) P2. Distal accessory cusp: absent (0); present (1). (14) P2. Occlusal shape: subrectangular (0); triangular (1). (15) P2. Buccal wall: rectilinear (0); conspicuous concavity (1). (16) P3. Robustness ratio [(maximum width/ maximum length) x 100]: slender P3 (less than 60) (0); Robust P3 (60 or more than 60) (1). (17) P3. Mesial accessory cusp: reduced or absent (0); present (1). (18) P3. Distal accessory cusp: present (0); absent (1). (19) P3. Buccal wall: rectilinear wall (0); conspicuous concavity (1). (20) P3. Basal cingulum: weak (0); strong (1). 4 (21) P3. Roots: 2 roots (0); 3 roots (1). (22) P3. Lingual bulge in the middle of the tooth: absent or reduced (0); Well- developed (1). (23) P4/3 length ratio. Maximum length of P4 in relation to maximum length of P3 ratio [(L P4/L P3) x 100]: less than 170 (0); more than 170 (1). (24) P4. Robustness ratio [(maximum width/ maximum length) x 100]: slender P4 (less than 60) (0); robust P4 (more than 60) (1). (25) P4 protocone: low (0); knoblike (1). (26) P4. Protocone cingulum: weak or absent (0); prominent and complete (1). (27) P4. Protocone: in front to the mesiobuccal corner (0), mesial to the mesiobuccal corner (1); distally displaced (2). (28) P4. Parastyle: Absent or weak (0); Strong and low (1); Strong and high (2). (29) P4. Buccal wall: rectilinear (0); with a pronounced concavity between paracone and metastyle (1). (30) P4. Basal cingulum surrounding the tooth: weak (0); strong (1). (31) Relative size between maximum width of M1 and the maximum length of P4: M1 larger than or equal in size to P4 (values >100) (0); M1 smaller than P4 (values between 70-100) (1); M1 much smaller than P4 (values <70) (2). (32) M1. Robustness ratio [(maximum width on the buccolingual area/ maximum length) x 100]: relatively robust M1 (from 130 to 150) (0); slender M1 (more than 150) (1), very slender M1 (more than 200) (2). (33) M1. Stylar area: small (0); enlarged (1). (34) M1. Metacone related to paracone: normal size (0); much reduced (1). (35) M1. Metaconule: present (0); absent (1). (36) M1. Protocone position: mesolingually located (0); almost centrally on the middle of the talon (1). (37) M1. Protocone shape: ridge-shaped (0); conical cusp-like shape (1). (38) M1. Lingual platform: not completely enclose the protocone (0); completely enclose the protocone (1). (39) M1. Length of the lingual wall: lesser than the buccal wall (unexpanded lingual platform) (0); longer than the buccal wall (moderately lingual platform) (1); much longer than the buccal wall (much expanded lingual platform) (2). 5 (40) M1. Lingual platform: oval shape (0); with a concavity in the middle point (1). (41) M1. Presence of a slender buccolingual crista from the paracone to the protocone: very reduced or absent (0); present (1). (42) p1. Present (0); absent (1). (43) p2. Position of the mesiodistal axis of p2: in line with the tooth row (0); buccolingually rotated (1). (44) p2. Distal accessory cuspid: absent (0); present (1). (45) p2. Basal cingulum: incomplete and weak (0); complete and strong (1). (46) p2. Mesial and distal cristids: weak cristids (0); thickened cristids (1). (47) p2. Robustness ratio [(maximum width/ maximum length) x 100]: slender p2 (less than 50) (0); relatively robust p2 (from 50 to 70) (1); very robust p2 (>70) (2). (48) p2 length compared to p3: p2 not reduced (0); p2 reduced (1). (49) p3. Mesial accessory cuspid: absent (0); present (1). (50) p3. Mesial and distal cristids: weak cristids (0); thickened cristids (1). (51) p3. Basal cingulum: incomplete and weak (0); complete and strong (1). (52) p3. Ratio distal thickened of p3 [(maximum width p3/ maximum length p3) x 100]: < 50, p3 elongated (0); from 50-70, p3 with distal expansion (1); > 70 distal part very thick (2). (53) p4. Length ratio in relation to m1 [(maximum length p4/ maximum length m1) x 100]: from 50 to 60, indicating a relatively not reduced p4 (0); more than 60, indicating a p4 relatively enlarged (1). (54) p4. Mesial accessory cuspid: absent or poorly developed (0); present, well developed (1). (55) p4. Distal accessory cuspid: High and well developed (0); low and reduced (1); absent (2). (56) p4. Mesial and distal cristids: weak cristids (0); thickened cristids (1). (57) p4. Basal cingulum: incomplete and weak (0); complete and strong (1). (58) p4. Buccal wall: straight (0); markedly concave (1). (59) p4. Backward inclination of the main cuspid: practically vertical (90º–80º) (0); with backward inclination (less than 80º) (1). (60) p4, unworn principal cusp: equals or exceeds height of m1 paraconid (0); lower than m1 paraconid (1). 6 (61) p4. Distal area: not buccolingually thickened, oval shaped (0); buccolingually thickened, quadrangular shaped (1); rhomboid shaped (2). (62) p4. Lingual expansion or basal bulge: absent (0); present (1). (63) m1. Height of protoconid in relation to paraconid: protoconid higher than paraconid (0); protoconid similar in height than the paraconid (1). (64) m1. Metaconid: Present (0); reduced/absent (1). (65) m1. Relative length of talonid with respect the total m1 length: the talonid 1/3 of the total length (0); equal or less than 1/4 of the total length (1). (66) m1. Width talonid ratio [(maximum talonid width/ maximum width in the base of the protoconid-metaconid) x 100]: talonid not reduced (values between 85-100) (0); reduced talonid (< 85) (1). (67) m1. Height of hypoconid: low (0); high (1). (68) m1. Position of hypoconid: labially located (0); centrally positioned or almost centrally positioned (1). (69) m1. Orientation of the hypoconid: almost vertical (0); orientated towards the protoconid (1). (70) m1. Entoconid: transform in an entocristid (0); absent (1) (71) m1 talonid: open and shallow basin with a reduced and low entocristid and a beveled lingual wall of the talonid (0); basin lost (1). (72) m1. Shape of the hypoconid: pyramidal (0); trending to a crest-like shape (1). (73) m1. Hypoconid size: medium (0); enlarged (1). (74) m1. Hypoconulid: very reduced to absent (0); present, not reduced (1). (75) m2. Always present (0); residual presence (present in a low rate <25% of the sample, comprising a reduced alveolus) or absent (1). (76) m2 paraconid: present (0); (1) very weak or absent (1). (77) m2. Metaconid: present (0); absent (1). Postcranial (78) Humerus. Delto-Pectoral crest most distal part: between the first third and the middle point of the diaphysis (0); exceeding half of the diaphysis (1). (79) Humerus. Development of the Deltoideus crest: Not very marked, normal development (0); large development, medially enlarged (1). 7 (80) Humerus. Development of the Pectoral crest: Not very marked, normal development (0); large development, laterally enlarged (1). (81) Humerus. Supraepicondylar crest: small size (0); enlarged and laterally projected (1). (82) Humerus. Medial epicondyle: small-sized, not proximodistally enlarged (0); large- sized, proximodistally enlarged and medially projected (1). (83) Humerus. Distal view of the distal epiphysis: Sub-quadrangular shape (0); rectangular shape, craniocaudally compressed (1). (84) Ulna. Proximal epiphysis: presence of two tubers (craniomedial and craniolateral one) on the olecranon (0); presence of one or even no tuber on the cranial part of the olecranon (1). (85) Ulna. Tuber olecrani: Rectilinear, scar for M. triceps brachii proximally projected (0); scar M. triceps brachii medially projected (1). (86) Ulna. Medial side of the most distal part of the diaphysis: small size and barely visible crest for the M. pronator quadratus, (0); moderate size of the crest for the M. pronator quadratus, occupying the last forth of the diaphysis (1); large development of this crest, medially projected, occupying the last third of the diaphysis (2). (87) Ulna. Articular circumference: robust, rostrally projected (0); reduced and integrated into the styloid process (1). (88) Ulna. Index of fossorial ability (IFA) according Rose et al. (2014) x 100: Values lesser than 22, = not fossorial (0); Values equal or higher than 22 = a relatively large olecranon process which is related with semifossorial traits in living badgers (1). (89) Radius. Proximal view of the proximal epiphysis: Oval- shaped (0); Subrectangular- shaped, craniocaudally reduced (1). (90) Radius. Radial tuberosity: Robust, oval, with the scar of the insertion of M. brachialis caudally projected (0); rounded scarf with a normal projection (1); barely marked scarf not caudally projected (2). (91) Radius. Medial side of the most distal part of the diaphysis: rectilinear wall, no visible crest or scarf for the M. pronator quadratus (0); large development of this crest, which occupies the last third of the diaphysis, and is medially projected, comprising a concave surface in the caudal side (1). 8 (92) Radius. Robustness ratio of the distal epiphysis [(maximum craniocaudal length/ maximum mediolateral length) x 100]: <70, subrectangular shaped (0); > 70 robust distal epiphysis, subquadrangular shape (1). (93) Radius. Distal epiphysis, medial styloid process: prominent, medially projected and relatively large (0); medium to small size, not medially projected (1). (94) Radius. Distal epiphysis, carpal articular surface: its medial point is relatively perpendicular to the main axis of the radius (0); its medial point conforms 45 degrees in relation to the main axis of the radius (proximally projected) (1). (95) Radius. Diaphysis: relatively rectilinear in medial view, non-craniocaudally widened (0); caudally curved at the distal part, comprising a concave surface (1). (96) Calcaneus. Calcaneal tuber: presence of two processes (medial and lateral) (0); presence of one process or even absence of processes (1). (97) Calcaneus. Calcaneal robustness index [(width of calcaneal head at widest point including sustentacular facet and the quadratum plantae process/ maximum length of the calnaneus) x 100]: values > 55, calcaneus relatively short (0); values between 55-45 normal length of the calcaneus (1); values < 45 calcaneus relatively elongated (2). (98) Calcaneus. Calcaneal tuber/calcaneal hell: slender (0); mediolaterally widen (1). (99) Calcaneus. Sustentacular facet: located close to the body of the calcaneus (0); medially projected (1). (100) Calcaneus. M. quadratus plantae process laterally expanded (0); M quadratus plantae process reduced and not projected (1). 9 Table S1. Geographic and temporal distribution of Eomellivora species, after Valenciano et al. (2015, 2017, 2019) and this paper. Species Age Locality and country Taxonomical references Age references Eomellivora sp. 11.62 Ma, late Aragonian (MN7+8) Hammerschmiede 5, Germany Kirscher et al. (2016) Kirscher et al. (2016) Eomellivora moralesi sp. nov. 11.21 Ma, late Aragonian (MN7+8) ACM/PTA-A2, Spain This paper This paper Eomellivora fricki Earliest Pannonian C, early Vallesian Gaiselberg, Austria Zapfe (1948); Valenciano et al. (2017) Valenciano et al. (2017) (MN9) Eomellivora fricki Pannonian D, early Vallesian (MN9) Wien XII-Altmannsdorf, Austria Pia (1939); Valenciano et al. (2017) Valenciano et al. (2017) Eomellivora fricki 9.76 Ma, early Vallesian (MN9) Can Llobateres 1, Spain Petter (1963); Crusafont-Pairó (1972); Valenciano Casanovas-Vilar et al. (2016) et al. (2019) Eomellivora fricki Early Vallesian (MN9) Los Valles de Fuentidueña, Spain Crusafont-Pairó & Ginsburg (1973); Ginsburg et al. Alberdi et al. (1981) (1981); this study Eomellivora piveteaui Vallesian (MN9–MN10) Yassiören, Turkey Ozansoy (1965); Wolsan & Semenov (1996); Kappelman et al. (2003) Valenciano et al. (2015) Eomellivora piveteaui Vallesian (MN9–MN10)? Wissberg, Germany Tobien (1955); Morlo (1997) Böhme et al. (2012); Pickford & Pourabrisham (2013) Eomellivora piveteaui Late Vallesian (MN10) Kalfa (=Calfa), Moldova Lungu (1978); Wolsan & Semenov (1996) Nesin and Nadachowsky (2001) Eomellivora piveteaui Late Vallesian (MN10) Buzhor 1 (= Bujor 1), Moldova Pevner et al. (2013) Nesin and Nadachowsky (2001) Eomellivora piveteaui 9.43–9.31 Ma, late Vallesian (MN10) Ravin de la Pluie, Greece Koufos (2012) Sen et al. (2000) Eomellivora piveteaui Subzone J2 (~9.6–9.3 Ma), late Vallesian Batallones-3, Spain Valenciano et al. (2015) Peláez-Campomanes et al. (MN10) (2017) Eomellivora piveteaui Subzone J2 (~9.6–9.3 Ma), late Vallesian Batallones-10, Spain Valenciano et al. (2015) Peláez-Campomanes et al. (MN10) (2017) Eomellivora sp. Early Turolian (MN11) Dorn-Dürkheim 1, Germany Morlo (1997) Franzen et al. (2013) Eomellivora wimani 9.79–9.76 Ma, Bahean Shangyingou (Sang-Yin-Kou, Lantian Basin Zdansky (1924); Wolsan & Semenov (1996) Kaakinen (2005); Zhang et al. locality 12), China (2013) 10 Eomellivora wimani 8.95 Ma, Bahean Liuwangou (Liu-Wan-Kou, Lantian Basin Zdansky (1924); Wolsan & Semenov (1996) Kaakinen (2005); Zhang et al. locality 31), China (2013) Eomellivora wimani Turolian (MN12?) Győrszentmárton 2 (=Pannonhalma), Kretzoi (1965); Wolsan & Semenov (1996) Gasparik (2001) Hungary Eomellivora wimani ~7.2–7.1 Ma, middle Turolian (MN12) Chimishliya (=Cimișlia), Moldova Simionescu (1938); Wolsan & Semenov (1996) Vangengeim and Tesakov (2008), reinterpreted; Vasiliev et al. (2011) Eomellivora wimani ~7.5–7.2 Ma, middle Turolian (MN12) Novaya Emetovka 2, Ukraine Orlov (1948); Wolsan & Semenov (1996) Vangengeim and Tesakov (2008), reinterpreted; Vasiliev et al. (2011) Eomellivora wimani Early Hemphillian (Hh2) Kern River Formation site 50, USA Stock & Hall (1933); Wolsan & Semenov (1996) Woodburne (2004); Janis et al. (2008) Eomellivora wimani Late Clarendonian (Cl3) North Tejon Hills Local Fauna, USA Baskin (1988) Woodburne (2004); Janis et al. (2008) Eomellivora wimani 8.2-7.1 Ma, Bahean Yuanmou-Xiahoe, China Zong (1994) Zhu et al. (2005); Dong & Qi (2013) Eomellivora cf. wimani 7.14-6.25 Ma, Baodean Lufeng hominoid locality, Shihuiba, China Qi (1985) Qi et al. (2006); Dong & Qi (2013) Eomellivora sp. Early Hemphillian (Hh1) Higgins Local Fauna (= Sebits Ranch) Dalquest and Patrick (1989) Janis et al. (2008) Eomellivora sp. Early Hemphillian (Hh1) Dove Spring Local Fauna Baskin (1988) Janis et al. (2008) Eomellivora sp. Early Hemphillian (Hh1) Smiths Valley Local Fauna Baskin (1988) Janis et al. (2008) Eomellivora ursugulo Early–middle Turolian (MN11–MN12) Grebeniki, Ukraine Orlov (1948); Wolsan & Semenov (1996) Vislobokova et al. (2001), Vangengeim and Tesakov (2008), reinterpreted; Vasiliev et al. (2011) Eomellivora ursugulo Middle–late Turolian (MN12–MN13) Taralik–Cher, Russia Lavrov & Gimranov (2018) Lavrov & Gimranov (2018) Eomellivora hungarica Late Turolian (MN13) Polgárdi 2, Hungary Kretzoi (1942); Wolsan & Semenov (1996) Freudenthal and Kordos (1989) Eomellivora sp. 11.15–11.06 Ma, early Vallesian (MN9) Gritsev (=Grytsiv), Ukraine Wolsan & Semenov (1996) Korotkevich et al. (1985); Kirscher et al. (2016) 11 Eomellivora sp. Early Vallesian (MN9) Borský Svätý Jur, Slovakia Lupták (1995) Sabol et al. (2004) Eomellivora sp. Early Turolian (MN11) Csákvár, Hungary Kretzoi (1942); Wolsan & Semenov (1996) Mein (1999) 12 Supplementary references Alberdi MT, López N, Morales J, Sesé C, Soria D. 1981. Bioestratigrafía y biogeografía de la fauna de mamíferos de los Valles de Fuentidueña (Segovia). Estud Geol. 37:503–511. Baskin JA. 1998. Mustelidae. In: Janis CM, Scott KM, Jacobs LL, editors. Evolution of Tertiary mammals of North America. Volume 1: Terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press; p. 152–173. Böhme M, Aiglstorfer M, Uhl D, Kullmer O. 2012. The antiquity of the Rhine River: stratigraphic coverage of the Dinotheriensande (Eppelsheim Fomation) of the Mainz Basin (Germany). PLoS ONE. 7:e36817. Casanovas-Vilar I, Garcés M, Van Dam J, García-Paredes I, Robles JM, Alba DM. 2016. An updated biostratigraphy for the late Aragonian and Vallesian of the Vallès-Penedès Basin (Catalonia). Geol Acta. 14:195–217. Crusafont-Pairó, M. 1972. Les Ischyrictis de la transition Vindobonien-Vallésien. Palaeovertebrata. 5:253–260. Crusafont-Pairó M, Ginsburg L. 1973. Les carnassiers fossiles de Los Vallès de Fuentiduena (Ségovie, Espagne). Bull Mus Natl Hist Nat. 131:29–45. Dalquest WW, Patrick DB. 1989. Small mammals from the early and medial Hemiphillian of Texas, with descriptions of a new bat and a gopher. J Vertebr Paleontol. 9:78–88. Dong W, Qi G-Q. 2013. Hominoid-producing localities and biostratigraphy in Yunnan. In: Wang X, Flynn LJ, Fortelius M, editors. Fossil mammals of Asia: Neogene biostratigraphy and chronology. New York: Columbia University Press, p. 1–25. Franzen JL, Pickford M, Costeur L. 2013. Palaeobiodiversity, palaeoecology, palaeobiogeography and biochronology of Dorn-Dürkheim 1—a summary. Palaeobiodiv Palaeoenviron. 93:277–284. Freudenthal M, Kordos L. 1989. Cricetus polgardensis sp. nov. and Cricetus kormosi Schaub, 1930 from the Late Miocene Polgárdi localities (Hungary). Scripta Geol. 89:71–100. Gasparik M. 2001. Neogene proboscidean remains from Hungary; an overview. Frag Palaeontol Hung. 19:61–77. Ginsburg L, Morales J, Soria D. 1981. Nuevos datos sobre los carnívoros de los Valles de Fuentidueña (Segovia). Estud Geol. 37:383–415. Koufos GD. 2012. New material of Carnivora (Mammalia) from the Late Miocene of Axios Valley, Macedonia, Greece. C R Palevol. 11:49–64. 13 Janis CM, Gunnell GF, Uhen MD, editors. 2008. Evolution of Tertiary Mammals of North America: Volume 2, Small Mammals, Xenarthrans, and Marine Mammals. Cambridge: Cambridge University Press. Kaakinen A. 2005. A long terrestrial sequence in Lantian - a window into the late Neogene palaeoenvironments of northern China [PhD dissertation]. Helsinki: University of Helsinki. Kappelman J, Duncan A, Feseha M, Lunkka J-P, Ekart D, Ryan TM, Swisher CC III. 2003. Chronology. In: Fortelius M, Kappelman J, Sen S, Bernor RL, editors. Geology and paleontology of the Miocene Sinap Formation, Turkey. New York: Columbia University Press; p. 41–66. Kirscher U, Prieto J, Bachtadse V, Aziz HA, Doppler G, Hagmaier M, Böhme M. 2016. A biochronologic tie-point for the base of the Tortonian stage in European terrestrial settings: Magnetostratigraphy of the topmost Upper Freshwater Molasse sediments of the North Alpine Foreland Basin in Bavaria (Germany). Newslett Stratigr. 49:445–467. Korotkevich E, Kushniruk V, Semenov Y, Chepaliga A. 1985. A new middle Sarmatian vertebrate fauna locality in Ukraine. Vestnik Zoologii. 29:81–82 (in Russian). Kretzoi M. 1942. Eomellivora von Polgárdi und Csákvár. Föld Közl. 72:318–323. Kretzoi M. 1965. Die Hipparion-Fauna von Győrszentmárton in NW-Ungarn. Ann hist-nat hungar. 57:127–143. Lavrov AV, Gimranov DO. 2018. First finding of a representative of giant mustelids of the genus Eomellivora (Carnivora, Mustelidae) in Russia (Tuva, Upper Miocene). Doklady Biol Sci. 480:82–84. Lungu AN. 1978. Hipparion Fauna of Middle Sarmatian of Moldova: Carnivores. Kishinev: Shtintsa (in Russian). Lupták P. 1995. First evidence of the Turolian carnivorous species Perunium ursogulo Orlov, 1948 (Mustelidae, Mammalia) from Slovakia. Slovak Geol Magaz. 2:171–174. Mein P. 1999. European Miocene mammal biochronology. In: Rössner GE, Heissig K, editors. The Miocene land mammals of Europe. Munich: Verlag Dr. Friedrich Pfiel; p. 25–38. Morlo M. 1997. Die Raubtiere (Mammalia, Carnivora) aus dem Turolium von Dorn-Dürkheim 1 (Rheinhessen). Teil 1: Mustelida [sic], Hyaenidae, Percrocutidae, Felidae. Cour Forsch-Inst Senck. 197:11–47. Nesin VA, Nadachowsky A. 2001. Late Miocene and Pliocene small mammal faunas (Insectivora, Lagomorpha, Rodentia) of southeastern Europe. Acta Zool Cracov. 44:107– 135. 14 Orlov YU. 1948. Perunium ursogulo Orlov, a new gigantic extinct mustelid (a contribution to the morphology of the skull and brain and to the phylogeny of Mustelidae). Acta Zool. 29:63–105. Ozansoy F. 1965. Étude des gisements continentaux et des mammifères du Cénozoïque de Turquie. Mém Soc Géol Fr. 44:1–92. Qi G. 1985. A preliminary report on Carnivora from the Ramapithecus locality, Lufeng, Yunnan. Acta Anthropol Sin. 4:33–43. Peláez-Campomanes P, Morales J, Álvarez Sierra MA, Hernández V. 2017. La edad de los yacimientos del Cerro de los Batallones. In: La colina de los tigres dientes de sable. Los yacimientos miocenos del Cerro de los Batallones (Torrejón de Velasco, Comunidad de Madrid). Madrid: Comunidad de Madrid; p. 147–161. Petter G. 1963. Contribution a l'étude des Mustélidés des bassins néogènes du Vallès-Pénédès et de Calatayud-Teruel. Mem Soc Géol Fr. 97:5–44. Pevner MA, Lungu AN, Vangengeym EA, Basilyan AE. 2013. Occurrences of Vallesian-age Hipparion fauna in Moldavia and their placement on the magnetic polarity scale. Int Geol Rev. 29:140–150. Pia J. 1939. Ein riesiger Honingsdachs (Mellivorine) aus dem Unterpliozän von Wien. Ann Naturhist Mus Wien. 50:537–583. Pickford M, Purabrishami Z. 2013. Deciphering Deinotheriensande deinotheriid diversity. Palaeobiodiv Palaeoenv. 93:121–150. Qi G,Dong W, Zheng L, Zhao L, Gao F, Yue L, Zhang Y. 2006. Taxonomy, age and environment status of the Yuanmou hominoids. Chin Sci Bull 51:704–712. Sabol M, Joniak P, Holec P. 2004. Succession(-s) of mammalian assemblages during the Neogene – A case study from the Slovak part of the Western Carpathians. Scripta Fac Sci Nat Univ Masaryk Brunensis. 31–32:65–84. Sen S, Koufos GD, Kondopoulou D, de Bonis L. 2000. Magnetostratigraphy of the late Miocene continental deposits of the lower Axios valley, Macedonia, Greece. Geol Soc Greece Spec Publ. 9:197–206. Simionescu I. 1938. Mamiferele pliocene de la Cimişlia (România). I Carnivore. Academia Română. Publicaţiunile fondului Vasile Adamachi. 9(50):1–30. Stock C, Hall ER. 1933. The Asiatic genus Eomellivora in the Pliocene of California. J Mammal. 14:63–65. 15 Tobien H. 1955. Neue und wenig bekannte Carnivoren aus den unterpliozänen Dinotheriensanden Rheinhessens. Notizbl Hess Landesamt Bodenforsch Wiesbaden. 83:7– 31. Valenciano A, Abella J, Sanisidro O, Harstone-Rose A, Álvarez-Sierra MÁ, Morales J. 2015. 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). J Vertebr Paleontol. 35:e934570. Valenciano A, Abella J, Göhlich UB, Ángeles Álvarez-Sierra M, Morales J. 2017. Re-evaluation of the very large Eomellivora fricki (Pia, 1939) (Carnivora, Mustelidae, Mellivorinae) from the late miocene of Austria. Palaeontol Electron. 20:17A. Valenciano A, Jiangzuo Q, Wang S, Li C, Zhang X, Ye J. 2019. First record of Hoplictis (Carnivora, Mustelidae) in East Asia from the Miocene of the Ulungur River Area, Xianjiang, Northwest China. Acta Geol Sin. 93:251–264. Valenciano A, Govender R. 2020. New fossils of Mellivora benfieldi (Mammalia, Carnivora, Mustelidae) from Langebaanweg, 'E' Quarry (South Africa, Early Pliocene): Re-evaluation of the African Neogene mellivorines. J Vertebr Paleontol. 40:e1817754. Vangengeim EA, Tesakov A. 2008. Maeotian mammalian localities of Eastern Paratethys: magnetochronology and positon in European continental scales. Stratigr Geol Correlation. 16:437–450. Vasiliev I, Iosifidi AG, Khramov AN, Krijgsman W, Kuiper K, Langereis CG, Popov VV, Stoica M, Tomsha VA, Yudin SV. 2011. Magnetostratigraphy and radio-isotope dating of upper Miocene–lower Pliocene sedimentary successions of the Black Sea Basin (Taman Peninsula, Russia). Palaeogeogr Palaeoclimatol Palaeoecol. 310:163–175.Vislobokova I, Sotnikova M, Dodonov A. 2001. Late Miocene - Pliocene mammalian faunas of Russia and neighbouring contries. Boll Soc Paleontol Ital. 40:307–313. Woodburne MO, editor. 2004. Late Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and Geochronology. New York: Columbia University Press. Woslan M, Semenov YA. 1996. A revision of the late Miocene mustelid carnivoran Eomellivora. Acta Zool Cracov. 39:593–604. Zapfe H. 1948. Neue Funde von Raubtieren aus dem Unterpliozän des Wiener Beckens. Sitz- Ber Öst Akad Wiss math-naturwiss Kl. 157:243–262. Zdansky O. 1924. Jungtertiäre carnivoren Chinas. Palaeontol Sin C. 2:1–149. Zhang Z-Q, Kaakinen A, Liu L-P, Lunkka JP, Sen S, Gose WA, Qiu Z-D, Zheng S-H, Fortelius M. 2013. Mammalian biochronology of the late Miocene Bahe Formation. In: Zhang X, Flynn LJ, 16 Fortelius M, editors. Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology. New York, N.Y: Columbia University Press; p. 187–202. Zhu R, Liu Q, Yao H, Guo Z, Deng C, Pan Y, Lü L, Chang Z, Gao F. 2005. Magnetostratigraphic dating of hominoid-bearing sediments at Zhupeng, Yuanmou Basin, southwestern China. Earth Planet Sci Lett 236:559–586. Zong G. 1994. On stratigraphic subdivision of hominoid fossil localities of Yuanmou, Yunnan. Hum Evol. 9:209–213. 17