This is the accepted version of the article: Valenciano, Alberto; Morales, Jorge; Azanza, Beatriz; [et al.]. «Aragonictis araid, gen. et sp. nov., a small-sized hypercarnivore (Carnivora, Mustelidae) from the late middle Miocene of the Iberian Peninsula (Spain)». Journal of vertebrate paleontology, (January 2022). DOI 10.1080/02724634.2021.2005615 This version is available at https://ddd.uab.cat/record/251463 under the terms of the license 1 Aragonictis araid gen. et sp. nov., a small-sized hypercarnivore (Carnivora, Mustelidae) 2 from the late middle Miocene of the Iberian Peninsula (Spain) 3 4 ALBERTO VALENCIANO,1,*, JORGE MORALES,2 BEATRIZ AZANZA,1 and 5 DANIEL DeMIGUEL3,1,4 6 1 7 Departamento de Ciencias de la Tierra, and Instituto Universitario de Investigación en 8 Ciencias Ambientales de Aragón (IUCA), Universidad de Zaragoza, 50009, Zaragoza, 9 Spain, a.valenciano@unizar.es; azanza@unizar.es; 2 10 Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC. C/ José 11 Gutiérrez Abascal, 2, 28006, Madrid, Spain. mcnm166@mncn.csic.es; 3 12 ARAID foundation, 50018, Zaragoza, Spain, demiguel@unizar.es; 4 13 Institut Català de Paleontologia Miquel Crusafont, Universidad Autònoma de 14 Barcelona, Campus de la UAB, 08193, Cerdanyola del Vallès, Barcelona, Spain. 15 16 *Corresponding author 17 18 RH: VALENCIANO ET AL.—A NEW SMALL MIDDLE MIOCENE MUSTELID 19 20 ABSTRACT—Small to medium-sized mustelids from the last 18 million years 21 represent a heterogeneous group of carnivorans with a wide-ranging record in the 22 northern hemisphere. They were first referred to the genera Mustela and Martes, but 23 lately ascribed to the latter, and hence considered as the longest-lived genus within 24 Mustelidae. However, a great many of these forms have been based upon fragmentary 25 material and Martes has conformed progressively to a wastebasket nomen for species of 1 26 uncertain relationships. Here, we describe dentognathic material of a small-sized 27 mustelid from three middle Miocene (MN7+8, latest Aragonian) localities of the Iberian 28 Peninsula that constitutes a new genus and species. Aragonictis araid gen. et sp. nov. 29 represents a distinct taxon if compared to early/middle Miocene forms ascribed to 30 “Martes” spp., especially the similar-sized early Miocene Circamustela? laevidens and 31 the middle Miocene “Martes” caedoti and “Martes” delphinensis. The finding of 32 particular features in A. araid (low p2-3, loss or reduction of the p4 accessory cuspid 33 with its main cuspid centrally located, presence of a sharp, beveled and lingually open 34 m1 talonid, and reduction of M1 lingual platform) indicates affinities with the late 35 Miocene Circamustela in the range of hypercarnivory. Our reassessment of “Martes” 36 indicates possible evidence of cladogenesis for Miocene mustelidae with, at least, two 37 different events being recognized in Europe—the latter during MN7+8 to MN9 with 38 presence of Aragonictis and Circamustela. The finding of A. araid further confirms the 39 presence of more densely forested environments than expected in inner Iberia during the 40 latest middle Miocene. 41 42 INTRODUCTION 43 44 Small to medium-sized mustelids from the early and middle Miocene are a 45 heterogeneous group of carnivorans with a wide-ranging record in Europe, Asia, and 46 North America (e.g., Baskin, 1998; Ginsburg, 1999; Peigné, 2012; Valenciano et al., 47 2020a). They were conventionally referred to both extant genera Mustela Linnaeus, 48 1758 (Mustelinae) and Martes Pinel, 1792 (Guloninae), albeit since the last century they 49 have been classified as Martes (Lartet, 1851; Depéret, 1887; Mayet, 1908; Hughes, 50 2012 and references herein). Accordingly, it represents the genus with the longest fossil 2 51 record in Mustelidae with more than 18 million years of duration from its emergence 52 (e.g., Dehm, 1950; Ginsburg, 1999; Peigné, 2012; Hughes, 2012). 53 However, the classification of a great many of these fossil forms has been based 54 upon fragmentary or incomplete material and have numerous plesiomorphic 55 traits/convergences with Martes, which makes of this genus a wastebasket nomen for 56 various small mustelids of uncertain relationships (Werdelin and Peigné, 2010; Samuels 57 et al., 2018). It remains to be discussed whether these Miocene forms should be 58 assigned to Martes (e.g., Jiangzuo et al., 2021), and emerges as an urgent task to 59 perform a comparison with the type species of the genus. While some of these forms 60 have morphological features closely related to Guloninae, others could instead represent 61 stem groups outside of the crown clade Guloninae (Anderson, 1970, 1994; Sato et al., 62 2003; Wang et al., 2012; Li et al., 2014; Samuels and Calvin, 2013; Samuels et al., 63 2018). For instance, Anderson (1994) proposed Martes laevidens Dehm, 1950 from the 64 early Miocene (MN3) of Wintershof-West (Germany) as the earliest member of the 65 genus, but such assignation to Martes has been recently discarded (Sato et al., 2003)— 66 though it is not fully discussed (as occur with the remaining European middle Miocene 67 species, Valenciano et al., 2020a). Within this complex scenario, it is debatable whether 68 the early late Miocene Martes melibulla Petter, 1963 from the Vallès-Penedès and 69 Madrid basins (Spain) represents the first true Martes (Valenciano, 2017; Valenciano et 70 al., 2020a), or it is instead the late Pliocene Martes wenzensis Stach, 1959 from Węże 1 71 (Poland) (Sato et al., 2003). Unfortunately, this systematics issue is far to be fully 72 clarified with the known mustelid material. 73 Herein, we report and describe new dentognathic remains of a small-sized 74 mustelid from three late middle Miocene Iberian localities (Aragonian, MN7+8, G3 75 local zone) (Fig. 1). This material greatly differs from that considered to date for Martes 3 76 and the other Miocene species included in the same genus. With the view to address this 77 challenge about this genus, the discovery of this new mustelid has the potential to shed 78 light on the obscure history and systematic intricacies of the genus Martes. 79 80 Geological Context and Age 81 We include material from the fossil sites of Andurriales and Toril 3A, which are 82 located in the Daroca area (province of Zaragoza, Spain) —one of the regions with the 83 highest concentration of mammalian fossil localities from Spain (Alcalá et al., 2000). 84 Both localities are magnetostratigraphically dated at 11.33 and 12.65 Ma, respectively, 85 hence belonging to local zone G3 (MN unit 7+8, upper Aragonian, middle Miocene) 86 (van Dam et al., 2014). The geology of the sites has been interpreted as an alluvial fan 87 that alternate with lake deposits (Alcalá et al., 2000; Azanza et al., 2004; van Dam et al., 88 2014; Barasoain and Azanza, 2018). Toril 3A exhibit an exceptional and abundant fauna 89 of vertebrates including amphibians, reptiles and birds, aquatic chelonians, small size 90 hornless ruminants, rhinos, cricetid species, and beavers, among others. This fauna 91 reveals the existence of areas of permanent or temporary fresh water and humid 92 conditions (Azanza et al., 2004; DeMiguel et al., 2011). The locality of Escobosa de 93 Calatañazor locality is a karstic infilling placed in the northern border of the Almazán 94 Basin (province of Soria, Spain). The micromammal association is correlated with local 95 zone G3 (as Nombrevilla 2), dated at 11.9 Ma (van Dam et al., 2014; Morales et al., 96 2018). 97 98 MATERIAL AND METHODS 99 100 Nomenclature and Measurements 4 101 Dental nomenclature follows Ginsburg (1999) and Smith and Dodson (2003). 102 Measurements were made using Mitutoyo Absolute digital caliper to the nearest 0.1 103 mm. 104 105 Abbreviations 106 Institutional Abbreviations—SNSB-BSPG, Staatliche Naturwissenschaftliche 107 Sammlungen Bayerns - Bayerische Staatssammlung für Paläontologie und Geologie, 108 Munich, Germany; FMNH, Field Museum of Natural History Chicago, Illinois, USA; 109 IPS, collection of the Institut Català de Paleontologia Miquel Crusafont (ICP), 110 Universitat Autònoma de Barcelona, Spain; MGUV, Museu de Geologia de la 111 Universitat de València, Burjassot; MNCN, Museo Nacional de Ciencias Naturales, 112 Madrid, Spain; MNHN, Muséum national d’Histoire naturelle, Paris, France; MPZ, 113 collection of the Museo de Ciencias Naturales de la Universidad de Zaragoza 114 (MCNUZ), Zaragoza, Spain; NMA, Naturmuseum Augsburg, Germany; NMB, 115 Naturhistorisches Museum Basel, Switzerland; PMU, Palaeontological Museum, 116 University of Uppsala, Uppsala, Sweeden; SMF, Senckenberg Forschungsinstitut und 117 Naturmuseum, Frankfurt, Germany; and UCBL-FSL, Université Claude Bernard Lyon 118 1 Lyon, France. 119 Other Abbreviations—c, lower canine; L, maximum length; M, upper molar; m, 120 lower molar; P, upper premolar; p, lower premolar; and W, maximum width. 121 122 Study Material 123 The study material includes unpublished dentognathic remains from the Spanish 124 localities of Andurriales and Escobosa de Calatañazor housed in the MNCN, and Toril 125 3A housed in the MCNUZ. The comparative material includes fossils of 5 126 “Martes”sanjoni (Mayet, 1908) and “Martes” burdigaliensis Beaumount, 1974 from 127 Artesilla (early Miocene, MN4, Spain) (Azanza et al., 1993) housed in the MCNUZ; 128 “Martes” sp., from La Barranca (middle Miocene, MN6, Spain) (Peigné et al., 2006) 129 housed in the MNCN; Martes ginsburgi Montoya, Morales and Abella, 2011 from 130 Venta del Moro (late Miocene, MN13, Spain) housed in the MGUV; “Martes” 131 sansaniensis (Lartet, 1851) from Sansan (middle Miocene, MN6, France) housed at 132 MNHN; Martes melibulla Petter, 1963 from the localities of Can Llobateres (late 133 Miocene, MN9, Spain) and Trinxera Nord de l’Autopista (late Miocene, MN10, Spain) 134 housed in the ICP; Martes aff. anderssoni from Can Poncic 1 (late Miocene, MN9) 135 (Petter, 1967) housed in the ICP; “Martes” munki Roger, 1900 from Sant Quirze del 136 Valles (middle Miocene, MN7+8, Spain) and Circamustela dechaseauxi Petter, 1967 137 from Can Llobateres housed in the ICP; Circamustela peignei Valenciano et al., 2020a 138 from Batallones-3 and 5 (late Miocene, MN10, Spain) housed in the MNCN; and both 139 Sinictis dolichognathus Zdansky, 1924 from Loc.108 of Zdansky (1924) (late Miocene 140 c. Turolian/Ventian, China), and Pekania palaeosinensis from localities 11, 30, 30 (5), 141 31, 108 and 111 of Zdansky (1924) (late Miocene c. Turolian/Ventian, China) housed in 142 the PMU. We also established comparisons with casts of “Martes” delphinensis 143 Depéret, 1892, “Martes” filholi (Depéret, 1887) and “M.” munki from La Grive (middle 144 Miocene, MN7+8, France) housed in the MNCN. Additionally, we examined 145 photographs of specimens of the following taxa: “M.” laevidens from Wintershof-West 146 (early Miocene, MN3, Germany) (Dehm, 1950) housed in the BSPG; five marten-like 147 taxa from Vieux-Collonges (middle Miocene, MN5, France) (“Martes” collengensis 148 Roth and Mein, 1987, “Martes” cadeoti Mein, 1958, “M.” burdigaliensis, “M.” munki 149 and “M.” filholi) housed in the UCBL; “M.” munki from Häder (middle Miocene, MN5, 6 150 Germany) (Roger, 1900) housed in the NMA; and Circamustela? from Dorn-Dürkheim 151 1 (late Miocene, MN11, Germany) (Morlo, 1997) housed in the SMF. 152 The extant specimens used for comparison are the following: Martes foina 153 (Erxleben, 1777) (MNCN); Martes martes Linnaeus, 1758 (MNCN); Pekania pennanti 154 (Erxleben, 1777) (FMNH); Mustela putorius Linnaeus, 1758 (MNCN); and Mustela 155 nivalis Linnaeus, 1766 (MNCN). 156 157 SYSTEMATIC PALEONTOLOGY 158 159 Order CARNIVORA Bowdich, 1821 160 Suborder CANIFORMIA Kretzoi, 1943 161 Family MUSTELIDAE Batsch, 1788 162 Subfamily GULONINAE Gray, 1825 163 Genus ARAGONICTIS gen. nov. 164 165 Type and Only Included Species—Aragonictis araid, gen. et sp. nov. 166 Etymology—Aragon-, for the autonomous community of Spain in where was 167 defined the Aragonian biozone (European Land Mammal Age: late early 168 Miocene/middle Miocene); -ictis, Greek suffix for weasel-like mammals. 169 Diagnosis—Mustelidae of small size. Slender dentition with narrow canines and 170 slender unicuspid premolars; p1 present; cuspids of p2-3 lower in height than the p4; 171 absence of diastema between p2-3; p4 long and unicuspid in the specimens from the 172 type locality, and with a very reduced distal accessory cuspid in the specimen from 173 Escobosa; m1 with subequal protoconid and paraconid, but slightly taller protoconid. 174 Small m1 metaconid. Sectorial and lingually open m1 talonid composed by a buccal 7 175 hypoconid relatively tall, beveled, lingually longer, with a reduced hypoconulid, and a 176 very low entocristid reaching the basal level of the paraconid crown; trapezoidal m2 177 with protoconid and metaconid of similar size, rectilinear buccal wall, and convex 178 lingual wall; slender P4 protocone mesially placed, paracone-metastyle blade elongated; 179 M1 mesiodistally narrow, with both mesial and distal walls straight but concave tooth 180 profile in mesial view; large parastylar area, reduced metacone, mesial crest-like 181 protocone, absent metaconule and short lingual platform. 182 183 ARAGONICTIS ARAID sp. nov. 184 (Figs. 2, 3) 185 Martes delphinensis Sesé 1980:133 186 187 Holotype—Left fragmentary hemimandible, MNCN 150.003, including the 188 complete dental formula c, p1-4, and m1-2. 189 Type Locality—Andurriales, MN7+8, local zone G3 (11.33 Ma; Van Dam et al., 190 2014), Zaragoza, Spain. 191 Other Localities—Toril 3A, MN7+8, local zone G3 (12.65 Ma; Van Dam et al., 192 2014), Zaragoza, Spain and Escobosa de Calatañazor, MN7+8, local zone G3 (ca. 12 193 Ma; Morales et al., 2018). 194 Referred Material—Andurriales: MNCN 150.004, right fragmentary 195 hemimandible with p2-3 and m1 (found associated with MNCN 150.003, probably 196 belonging to the same individual); Toril 3A: MPZ 2020/474, left dp4; MPZ 2020/482, 197 left fragmentary M1; Escobosa de Calatañazor: MNCN 26577, right fragmentary 198 hemimandible with p2, p4-m2 and MNCN 150.007, left fragmentary maxillary with P4- 199 M1. 8 200 Etymology—After ARAID (acronym for the Fundación Agencia Aragonesa para 201 la Investigación y el Desarrollo—in English, Aragonese Foundation for Research and 202 Development), an agency created by the Government of Aragon with the aim of 203 promoting research, development and innovation. In recognition of its implication in the 204 research of vertebrate paleontology in Aragon. 205 Diagnosis—As for genus. 206 Description—The mandibular corpus is long and low (Fig. 2). Two mental 207 foramina are present, one below the p2 and other at the level of the p3. The c is narrow. 208 The lower premolars are elongated and unicuspids. There is not diastema between the 209 lower premolars (Fig. 2A-C). The p1 is present. The cuspid of both p2-3 are lower than 210 p4 and placed mesially. The p4 is tall with a cuspid situated in the middle of the tooth 211 and reaches the high of the m1 paraconid. A low and reduced distal accessory cusp is 212 present in the specimen MNCN 26577 (Fig. 2M-O). A mesial and distal crown 213 cingulids are present. The m1 is long compared with the p4. The trigonid represents 214 three fourths of the total length of the molar. The protoconid is the tallest cuspid, 215 although the paraconid is relatively tall. It has a small metaconid, and a bucco-lingually 216 reduced talonid. The hypoconid is clearly separated from the postprotocristid. It is 217 buccally placed, and shows a beveled morphology, being bulbous in the buccal wall, 218 and with a marked lingual slope in the lingual wall towards the entoconid crista. There 219 are no entoconid or hypoconulid in the type, but a small hypoconulid is presents in 220 MNCN 26577. The m1 entocristid is very low reaching the basal level of the paraconid 221 crown, providing an open talonid valley (Fig. 2B-E-O). This area is broken in MNCN 222 26577 (Fig. 2M-O). The m2 is trapezoidal. It has a protoconid and a metaconid of 223 similar size. The trigonid is larger than the talonid. The buccal wall is rectilinear and 224 includes the paraconid, protoconid and hypoconid. On the contrary, the lingual wall is 9 225 convex. The lower deciduous carnassial (dp4) (Fig. 2J-L) is very similar to that of the 226 described m1. It differs in an acuter talonid, being relatively shorter and sharper with a 227 smaller hypoconid. The P4 paracone-metastyle blade is elongated, compared with the 228 M1 width (Fig. 3). The parastyle is very low and the protocone is mesially located and 229 isolated. The M1 lacks the typical eight-shaped mustelid morphology and is reduced 230 compared with extant gulonines (Fig. 3). The mesial and distal walls are rectilinear, 231 although the tooth profile is concave in mesial view. It has a large parastylar area, and a 232 reduced metacone compared to the paracone. There is not metaconule. The protocone is 233 crest-like and mesially placed. The lingual platform is not mesiodistally expanded. The 234 specimen MPZ 2020/482 is larger to MNCN 150.007, but morphologically similar, 235 including a crest-like and mesially placed (Fig. 3B). 236 Measurements—Measurements are provided in Table 1. 237 238 DISCUSSION 239 240 The new mustelid here reported from the latest middle Miocene of the Iberian 241 Peninsula differs from the living genera Mustela and Martes in multiple morphological 242 traits. It differs from Mustela (type species Mustela erminea Linnaeus, 1758), as well as 243 other living Mustela spp., in a larger size, longer and slender mandible, presence of p1, 244 longer lower premolars (especially the p4), taller m1 protocone, presence of m1 245 metaconid, larger and wider m1 talonid with smaller hypoconid, and well-developed 246 entocristid. M1 with both larger parastylar area and metacone, along with absence of 247 narrowing of the crown at about mid-width. 248 The seven extant species of marten sensu stricto (Martes) are categorized among 249 other traits by having long, relatively narrow skulls, a long auditory bulla, a large P4 250 protocone (less developed in type species of the genus M. foina), an enlarged M1 10 251 lingual platform (inner lobe), and a relatively elongated and basined m1 talonid, with a 252 conical hypoconid linking the metaconid by an entocristid (Anderson, 1970, 1994; 253 Jiangzuo et al., 2021). The new Spanish mustelid also differs from M. foina and other 254 living Martes spp. in a smaller size (Figs. 4-5), unicuspid and slenderer p4 comprising a 255 centrally located cuspid, m1 without mesoconid, stronger metaconid, slenderer talonid, 256 with much lower entocristid, and more open talonid valley, plus an absent or reduced 257 hypoconulid. It also has a lesser subquadrangular talonid in occlusal view. The m2 is 258 more asymmetric, and comprises a more developed talonid. The P4 protocone is 259 slenderer and the M1 has a larger parastylar area, an absence of both metaconule and 260 narrowing of the crown at about mid-width, besides of a much more reduced lingual 261 platform. Therefore, this new form can be confidently described as a genus distinct of 262 Martes. However, based on the hitherto known fossils of this new form, it is difficult to 263 determine with accuracy its systematic position at the subfamily level. Accordingly, we 264 provisionally place it into the extant subfamily Guloninae, whose early members, as 265 Dehmictis Ginsburg and Morales, 1992, and Iberictis Ginsburg and Morales, 1992, are 266 recorded in the early Miocene of Central and Western Europe (Valenciano et al., 267 2020b). 268 The described specimens of Aragonictis araid shown a range of variability in 269 dental size and morphology comparable to those of marten-like mustelids found in the 270 localities of Wintershof-West (Dehm, 1950) and Sansan (Peigné, 2012), being also 271 similar to living Martes foina (Figs. 4-5). Aragonictis araid specimens from Escobosa 272 are easily recognized by being the smaller ones, whereas specimens from Toril 3A 273 represent the largest in size among the sample (Figs. 4-5, Table 1). However, the fact of 274 sharing a lower p2, p4 with the main cuspid centrally located, together with a great 275 reduction of the p4 accessory cuspid, a similar m1 talonid (lingually broken in the one 11 276 from Escobosa), and a reduced M1 lingual platform with a crest-like protocone, allows 277 the assignment of all these fossils into the same taxon. 278 Over the course of the 19th and 20th centuries, fourteen species of Martes have 279 been described in the Miocene of Europe, ranging from MN3 to MN13 (Ginsburg, 280 1999; Montoya et al., 2011; Peigné, 2012; Hughes, 2012). They represent a group 281 widely heterogeneous and unevenly known, which has led researchers to often place 282 their mustelid material of small size and marten-like morphology inside of the 283 wastebasket name of Martes. Sato et al. (2003) specified Martes wenzensis from the late 284 Pliocene of Węże 1 (Poland) as the oldest known Martes. This taxon is very similar in 285 morphology to living Martes martes, but larger in size (Anderson, 1970), and its 286 assignment to the genus is not questionable. Despite this, Martes remains can be found 287 in older sediments than late Pliocene. Several key features of marten are already present 288 in late Miocene forms such as Martes melibulla from the Vallès-Penedès localities of 289 Can Llobateres (late Miocene, MN9) and Trinxera Nord de l’Autopista (late Miocene, 290 MN10) (Petter, 1963; Pons-Moyà, 1990), as well as the early late Miocene Martes aff. 291 anderssoni of Can Poncic 1 (MN9), also in the Vallès-Penedès Basin (Petter, 1967). 292 These traits are the enlargement of the M1 lingual platform, and a relatively elongated 293 and basined m1 talonid with a conical hypoconid connected with the metaconid by an 294 entocristid. More complete material of these forms would be needed to further 295 investigate the validity of this hypothesis. 296 Independent of which taxa represent the oldest record of Martes, there are 297 findings that clearly indicate that abundant small marten-like mustelids from the early 298 and middle Miocene of Eurasia ascribed to “Martes” show a dissimilar morphology 299 with Martes and stand in need of thorough revision (e.g., Hughes, 2012; Peigné, 2012). 300 Among these heterogeneous sample, four distinct groups can be distinguished: 12 301 (1) “Martes” spp. that are close to Martes but exhibit primitive traits which 302 distinguished them from this genus. This group is composed by the small-sized “M.” 303 cadeoti (Fig. 6A-D) and “M.” collongensis from Vieux-Collonges (middle Miocene, 304 MN5, France) (Mein, 1958; Roth and Mein, 1987), “M.” burdigaliensis from Vieux- 305 Collonges, Artesilla and Artenay (early and middle Miocene, MN4-MN5) (Beaumount, 306 1974; Azanza et al., 1993; Ginsburg, 2002), and by the large-sized “M.” munki from 307 Häder, Sandelzhausen, Sant Quirze del Vallès, La Grive (middle Miocene, MN5- 308 MN7+8) (Roger, 1900; Villalta Comella and Crusafont Pairó 1943; Viret 1951; Mein, 309 1958; Nagel et al., 2009) and “Martes” sansaniensis from Sansan (middle Miocene, 310 MN6) (Peigné, 2012). All the above include taxa that resemble Martes in having a 311 similar carnassial and M1; a p4 distal accessory cuspid; a relatively elongated and 312 basined m1 talonid (shorter in “M.” burdigalensis) with a conical hypoconid linking the 313 metaconid by an entocristid; a large P4 protocone; an enlarged M1 lingual platform; and 314 presence of a narrow M1 crown at about mid-width (Fig. 6A-D). Furthermore, all of 315 these extinct forms share several primitive traits: diastema between p2-3, presence of a 316 m1 protoconid very tall, m1 metaconid well developed and tall (especially in “M.” 317 cadeoti, and “M.” collongensis), M1 with distinct proportions (being more elongated 318 buccolingually, Fig. 5) with a M1 parastyle and metacone more developed, a 319 metaconule, and a protocone located more buccally compared with living Martes. 320 Aragonictis araid is different from this group having a lower p2-3; absence of diastema 321 in the lower dentition; absence or reduction of the distal accessory p4; m1 paraconid and 322 protoconid close in high, m1 talonid mesiodistally and buccolingually shorter with a 323 beveled hypoconid, a lower entocristid, reduced hypoconulid and distal cingulid, and a 324 more open valley; reduced m2; slenderer P4 protocone; and M1 with larger parastyle 13 325 area, reduced metacone, absent metaconule, absence of narrowing of the crown at about 326 mid-width, and a much reduced lingual platform. 327 (2) “Martes” sainjoni from Artenay (type locality), Artesilla and Bezian (Mayet, 328 1908; Ginsburg and Bulot, 1982; Azanza et al., 1993; Ginsburg, 2002), for which only 329 the lower dentition is known. This early-middle Miocene mustelid is different from 330 Aragonictis and Martes in being a more robust form, and having a distinct m1talonid 331 including a massive hypoconid, centrally positioned, and a strong cingulum. More 332 specifically, A. araid differs from “M.” sainjoni in a smaller size; slenderer mandibular 333 corpus; lower p4; absence of cingulid in both p4 and m1; shorter length of the trigonid 334 and less conical m1 hypoconid. Thus, “M.” sainjoni emerges as closer to basal Miocene 335 gulonines such as Dehmictis Ginsburg and Morales, 1992, Laphictis Viret, 1933, and 336 Ischyrictis Helbing, 1930. 337 (3) “Martes” filholi from La Grive (type locality), Viex-Collonges (Depéret, 1887; 338 Viret, 1951; Mein, 1958), and (albeit with certain doubts) Rudabánya (Werdelin, 2005). 339 It has diastema in the lower dentition; higher p2-3; longer m1 talonid; much developed 340 m2; shorter P4; and a much more distinct M1 with reduced parastyle, enlarged 341 metacone, a much-enlarged lingual platform and a very marked narrowing of the crown 342 at about mid-width than A. araid. It is clearly different from Martes, being probably 343 more related to Trochictis Meyer, 1842, or related genera (see Morlo et al., 2021). 344 (4) Small sized-mustelids with slender dentition comprising “Martes” 345 delphinensis and “Martes” laevidens. The former, which conforms the smallest mustelid 346 among the analyzed sample (Fig. 4), is recorded throughout the middle Miocene of 347 Western Europe, in the fossil sites of La Grive (type locality), Vieux-Collonges, 348 Manchones (MN6, Calatayud-Daroca Basin, Spain), Hostalets de Pierola (MN7+8, 349 Vallès-Penedès Basin, Spain) (Depéret, 1892; Villalta Comella and Crusafont Pairó 14 350 1943; Viret, 1951; Mein, 1958; Petter, 1976), and in all probability La Barranca (MN6, 351 Calatayud-Daroca Basin, Spain) (described previously as Martes sp. by Peigné et al., 352 2006). “Martes” delphinensis is represented only by mandibles and lower dentition, so 353 it has been scarcely studied in the past. Nevertheless, Mein (1958) described and figured 354 a maxillary with a complete M1 (Mein, 1958;fig. 56) for which a re-examination allows 355 us to confirm that it may belong to the stem mephitid Proputorius Filhol, 1890 which 356 also occurs in the locality (Proputorius pusillus [Viret, 1951]). The dimensions of A. 357 araid and its superficial similarities with the small-size mustelid “M.” delphinensis 358 makes its comparison somewhat intricate. Indeed, the mandible MNCN 26577 from 359 Escobosa was previously described as “M.” delphinensis in Sesé (1980). Aragonictis 360 araid differs from “M.” delphinensis in a larger size; a lower p2-3; a central position of 361 the main cuspid of the p4; an absence/reduction of the p4 distal accessory cuspid; a m1 362 paraconid higher; and a more open and longer m1 talonid with a much lower entocristid. 363 With regard to “Martes” laevidens from the early Miocene (MN3) of Wintershof-West 364 (Germany) (Dehm, 1950), this can be considered as the oldest species in the group. Sato 365 et al. (2003) discarded its assignation to Martes based on the basiocranial fragment 366 described by Dehm (1950:fig. 130), but they did not analyze its dentition. The sample 367 described in Dehm (1950) has some size variability (Figs. 4-5). The lower dentition is 368 slender, and has a diastema between p2-3, a well-developed distal accessory cuspid in 369 p4, and a buccolingually reduced m1 talonid. The P4 protocone is relatively robust 370 when compared with A. araid, and the M1 (although it shows some variability, see Fig. 371 6E-N) is characterized by an elongated parastylar area, a reduced metacone (almost 372 absent in SNSB-BSPG 1937 II 13372, see Fig. 6L), and a reduced lingual platform with 373 some degree of expansion. It must be emphasized that the original ascription by Dehm 374 (1950) of the M1 SNSB-BSPG 1937 II 13723 (Fig. 6O) with mustelid features to “M.” 15 375 laevidens, must be discarded due to its distinct morphology. Aragonictis araid differs 376 from “M.” laevidens in having a less sigmoid dp4 with a less developed metaconid; 377 absence of both diastema between p2-3 and lower cuspids; p4 with less crowned mesial 378 and distal cingulids; absence or reduction of p4 distal accessory cuspid; beveled m1 379 hypoconid, slenderer P4; as well as smaller M1 parastylar area with more developed M1 380 metacone. The extremely reduced metacone in the M1 SNSB-BSPG 1937 II 13372 381 (Fig. 6L) of “M.” laevidens is almost fuses in a crista with the paracone (Schmidt- 382 Kittler, 1981). This feature is shared with Circamustela dechaseauxi (type specie of 383 Circamustela Petter, 1967) from Can Llobateres (late Miocene, MN9, Vallès-Penedès 384 Basin). The m1 of both species—which is the only possible additional comparison— 385 shows the same morphological pattern in which the metaconid is reduced and the 386 talonid is narrow (Petter, 1967, 1976; Valenciano et al., 2020a). For all these reasons, 387 and considering the limitation of comparisons in the rest of the dentition and the 388 important separation in age between these two species, it seems convenient to reclassify 389 the Wintershof-West species as Circamustela? laevidens (Dehm, 1950). 390 Additionally, albeit A. araid shares several dental traits (including a slender 391 dentition, a similar P4, and a reduced M1 lingual platform; Valenciano et al., 2020a) 392 with the early late Vallesian Circamustela spp., it is different in many others. For 393 instance, the new species is smaller in size, and it has absence of diastema between p2- 394 3, as well as lower p2-3, presence of m1 metaconid (quite reduced in C. dechaseauxi), 395 slenderer m1 talonid with lower entocristid and more open valley, M1 more concave in 396 distal view with deeper trigone valley, and more developed metacone than the type 397 specie. Moreover, a similar sized M1 of Circamustela? appeared in younger (early 398 Turolian, MN11) sediments of Dorn-Dürkheim 1 (Germany) (Morlo, 1997). Despite its 399 small size, (closer to A. araid than to Circamustela spp., Fig. 5), its reduced parastylar 16 400 area, and the narrowing of the crown at about mid-width, as well as the less reduced 401 lingual platform, allows us to propose that the German M1 is closer related to 402 Circamustela than to Aragonictis. 403 An overview of the described groups suggests the inclusion of Aragonictis within 404 the “M.” delphinensis and C.? laevidens group, being probably related to Circamustela. 405 However, more dentognathic material is necessary in order to further determine its 406 systematic affinities and verify whether these similarities are a consequence of a 407 convergence towards hypercarnivorism or are instead result of close evolutionary 408 relationships 409 410 Paleobiological Insights 411 Compared with small to medium-sized living gulonines (e.g., stone marten, beech 412 marten, fisher) of omnivore diets (Larivière and Jennings, 2009) and with Miocene 413 forms of “Martes” from Europe (Fig. 4), A. araid shares similarly narrower teeth (e.g., 414 m1, P4 and M1) to those of the early Miocene Circamustela? laevidens and the late 415 Miocene Circamustela (Vallesian, MN9-10) and Sinictis (Turolian, MN11-MN13). 416 Because these late Miocene mustelids have been customarily interpreted as 417 hypercarnivores (Valenciano et al., 2020a), we interpret A. araid as a hypercarnivore 418 form as well. 419 It is remarkable how the discovery of this new Iberian mustelid and our review of 420 Miocene “Martes” suggest the occurrence of several cladogenesis events in Central and 421 Western Europe in this epoch. A first event took place at the end of the early Miocene 422 (MN3-4) and the beginning of the middle Miocene (MN5) in both Central (Germany) 423 and Western (France and Iberian Peninsula) Europe. During such a relatively long 424 event, roughly coinciding with the Miocene Climatic Optimum (Zachos et al., 2008), 17 425 members of the group of “Martes” appeared. An increase in the carnivoran´s Spanish 426 turnover rate during the late Aragonian MN7+8, as noticed by Morales et al. (2015), 427 informs about a second cladogenesis event. It was probably triggered, among other 428 factors, by an increase of the precipitation levels compared to the previous biozone (Van 429 Dam, 2006), thus increasing the forest areas of the region. The presence of small 430 mustelids has been traditionally considered as indicative of forested habitat (Ginsburg 431 and Morales, 1992; Baskin, 1998). Therefore, the large diversity of small mustelids in 432 Toril 3A, which comprises several species of “Martes” and the new form A. araid, 433 might suggest the presence of forest patches embedded in a mosaic environment for this 434 locality at ca. 12.65 Ma. This fits well with the hypothesis of a climate change towards 435 more humid conditions and increased canopy cover by the latest Aragonian in the 436 Daroca-Nombrevilla area (DeMiguel et al., 2011, 2012; García Paredes et al., 2016). 437 Such kind of habitats would have probably been suitable for other contemporaneous 438 mammals from the same basin customarily interpreted as forest-dwellers. Therefore, the 439 discovery of A. araid strongly indicates that a cladogenesis event for small to medium- 440 sized mustelids of hypercarnivorous dentition took place, at least at a regional scale, at 441 the end of the Aragonian (ca. 12.65 Ma). As such, some of these clades seem to have 442 increased in size during the late Miocene, as occurred for the Vallesian Circamustela 443 (Vallès-Penedès and Tagus Basins, Spain) (Valenciano et al., 2020a), and for later 444 forms such as the Turolian Sinictis (China and Greece) (Zdansky, 1924; Valenciano et 445 al., 2020a). An in-depth review of marten and weasel-like forms from other 446 contemporaneous localities from Eurasia would be the logical next step to further 447 investigate this cladogenesis event at a more global scale during the latest Aragonian. 448 449 CONCLUSIONS 18 450 451 We describe new mustelid material from the latest middle Miocene sites of 452 Andurriales, Toril 3A and Escobosa de Calatañazor (Iberian Peninsula). This form 453 constitutes a new genus and species that we name as Aragonictis araid. 454 Our analyses show that A. araid is morphologically different from living Mustela 455 and Martes, and also differs from the early and middle European Miocene forms of 456 Europe ascribed sensu lato to “Martes” spp., thus rejecting the conventional view that 457 all small to medium-sized Miocene mustelids belong to Martes. A preliminary 458 reassessment of these heterogenous taxa (formerly considered as Martes) allows us to 459 distinguish four different groups: (1) early to middle Miocene “Martes” spp.—distinct 460 from Martes sensu stricto—including “M.” collongensis, “M.” cadeoti, “M.” 461 burdigaliensis, “M.” munki and “M.” sansaniensis; (2) early to middle Miocene 462 “Martes” sainjoni from Western Europe—much closer to basal Miocene gulonine (such 463 as Dehmictis, Laphictis, and Ischyrictis) than to Martes; (3) middle Miocene “Martes” 464 filholi—a form probably related with Trochictis; and (4) early Miocene “Martes 465 delphinensis” and “Martes” laevidens—the latter being herein reassessed as 466 Circamustela?. 467 Finally, from a paleobiological viewpoint, we interpreted the new Iberian 468 Aragonictis araid as a hypercarnivorous mustelid that lived in forest patches of mosaic 469 environments. 470 471 ACKNOWLEDGMENTS 472 473 We thank the following colleagues for kindly providing us comparative material 474 and photographs of fossil mustelids: P. Mein and E. Robert (UCBL, France), C. Argot 475 (MNHN), G. Rössner and M. Schellenberger (BSPG, Germany), M. Morlo, O. Kullmer 19 476 and C. Hemm (SMF, Germany), M. Rummel (Naturmuseum Augsburg, Germany), S. 477 Fraile and A. Garvía (MNCN, Spain), J. M. Robles and D. M. Alba (ICP, Spain), I. 478 Canudo (MCNUZ, Spain), L. Costeur (NMB, Switzerland), J. O. R. Ebbestad and V. 479 Berg-Madsen (PMU, Sweeden), and the staff of Servicio de Técnicas No Destructivas: 480 Microscopía Electrónica y Confocal y Espectroscopía del MNCN-CSIC. We also thank 481 C. Langa for finding the fossils from Andurriales, and E. Cantero (MNCN) for the 482 preparation of the material. We are indebted to the editor T. Martin and the reviewers L. 483 Werdelin and M. Morlo for their useful comments and suggestions, which made 484 meaningful improvements to the original manuscript. This study was supported by the 485 Government of Aragon (Group ref. E33_20R), the Research Group UCM 910607, the 486 Spanish Ministry of Economy and Competitiveness and FEDER funds (Research 487 Projects PGC2018-094122-B-100 and PID2020-116220GB-I00), and the Spanish 488 Ministry of Science, Innovation, and Universities (“Juan de la Cierva Formación”, ref. 489 FJC2018-036669-I). 490 491 LITERATURE CITED 492 493 Alcalá, L., A. M. Alonso-Zarza, M. A. Álvarez, B. Azanza, J. P. Calvo, J. C. Cañaveras, 494 J.A. van Dam, M. Garcés, W. Krijgsman, A. J. van der Meulen, J. Morales, P. 495 Peláez, A. Pérez-González, S. Sánchez, R. Sancho, and E. Sanz. 2000. El registro 496 sedimentario y faunístico de las cuencas de Calatayud-Daroca y Teruel. Evolución 497 paleoambiental y paleoclimática durante el Neógeno. Revista de la Sociedad 498 Geológica de España13:323–343. 499 Anderson, E. 1994. Evolution, prehistoric distribution and systematics of Martes; 500 pp.13–25 in S. W. Buskirk, A. S. Harestad, Raphael and R. A. Powell (eds.), 20 501 Martens, Sables and Fishers: Biology and Conservation. Cornell University Press, 502 Ithaca, New York. 503 Anderson, E. 1970. Quaternary evolution of the genus Martes (Carnivora, Mustelidae). 504 Acta Zoologica Fennica 130:1–132. 505 Azanza, B., E. Cerdeño, L. Ginsburg, J. Van der Made, J. Morales, and P. Tassy. 1993. 506 Les grands mammifères du Miocène inférieur d’Artesilla, bassin de Calatayud- 507 Teruel (province de Saragosse, Espagne). Bulletin du Museum National d'Histoire 508 naturelle 15:105–153. 509 Azanza, B., A. M. Alonso-Zarza, M. A. Álvarez-Sierra, J. P. Calvo, S. Fraile, I. García- 510 Paredes, E. Gómez, M. Hernández, A. van der Meulen, D. DeMiguel, P. Montoya, J. 511 Morales, X. Murelaga, P. Peláez-Campomanes, B. Pérez, V. Quiralte, M. J. Salesa, I. 512 M., Sánchez, A. Sánchez, and D. Soria. 2004. Los yacimientos de vertebrados 513 continentales del Aragoniense superior (Mioceno medio) de Toril, Cuenca de 514 Calatayud-Daroca. Geo-Temas 6:271–274. 515 Barasoain, D. and B. Azanza. 2018. Geoheritage and education: a practical example 516 from the Rhinoceros of Toril 3 (Calatayud-Daroca Basin, Spain). Geoheritage 517 10:363–374. 518 Baskin, J. A. 1998. Mustelidae; pp. 152-173 in C. M. Janis, K. M. Scott, L. L. Jacobs 519 (eds.), Evolution of Tertiary Mammals of North America, Volume 1: Terrestrial 520 Carnivores, Ungulates, and Ungulate-like Mammals. Cambridge University Press, 521 Cambridge. 522 Batsch, A. J. G. C. 1788. Versuch einer Anleitung, zur Kenntniβ und Geschichte der 523 Thiere und Mineralien, für akademische Vorlesungen entworfen und mit den 524 nöthigsten Abbildungen versehen. Erster Theil. Academische Buchhandlung, Jena. 21 525 Beaumount, G. de 1974. Un nouveau Mustélidé (Carnivora), Martes burdigaliensis n. 526 sp., du Miocène inférieur de Vieux-Collonges, Rhône. Comptes Rendus des Séances, 527 Société de Physique et d’Histoire naturelle de Genève 9:81–83. 528 Bowdich, T. E. 1821. An Analysis of the Natural Classifications of Mammalia For The 529 Use Of Students And Travelers. J. Smith, Paris, 115 pp. 530 Dehm, R. 1950. Die Raubtiere aus dem Mittel-Miocän (Burdigalium) von Wintershof- 531 West bei Eichstätt in Bayern. Abhandlungen der Bayerischen Akademie der 532 Wissenschaften. Mathematisch-naturwissenschaftliche Klasse, n. F. 58: 1-158. 533 DeMiguel, D., B. Azanza, and J. Morales. 2011. Paleoenvironments and paleoclimate of 534 the Middle Miocene of central Spain: a reconstruction from dental wear of 535 ruminants. Palaeogeography, Palaeoclimatology, Palaeoecology 302:452–463. 536 DeMiguel, D., I. M. Sánchez, D. M. Alba, J. Galindo, J. M. Robles, and S. Moyà-Solà. 537 2012. First evidence of Tethytragus Azanza and Morales, 1994 (Ruminantia, 538 Bovidae), in the Miocene of the Vallès-Penedès Basin (Spain). Journal of Vertebrate 539 Paleontology 32:1457–1462. 540 Depéret, C. 1887. Recherches sur la succession des faunes de vertébrés miocènes de la 541 vallée du Rhône. Archives du Muséum d’Histoire naturelle de Lyon 4:45–269. 542 Depéret, C. 1892. La faune des Mammifères miocènes de la Grive-Saint-Alban (Isère) 543 et de quelques autres localités du bassin du Rhône. Archives du Muséum d’Histoire 544 naturelle de Lyon 5:1–93. 545 Erxleben, J. C. P. 1777. Systema regni animalis per classes, ordines, genera, species, 546 varietates cum synonymia et historia animalium. Classis I. Mammalia. Weigand, 547 Leipzig: I-XLVII:1–636. 548 Filhol, H. 1890. Étude sur les mammifères de Sansan. Bibliothèque des Hautes Études 549 section des Sciences naturelles 37:1–319. 22 550 García-Paredes, I., M. A. Álvarez-Sierra, L. W. Van den Hoek Ostende, V. Hernández- 551 Ballarín, K. Hordijk, P. López-Guerrero, A. Oliver, and P. Peláez-Campomanes. 552 2016. The Aragonian and Vallesian high-resolution micromammal succession from 553 the Calatayud-Montalbán Basin (Aragón, Spain). Comptes Rendus Palevol 15:781– 554 789. 555 Ginsburg, L. 1999. Order Carnivora; pp. 109–148 in G. E. Rössner, and K. Heissig 556 (eds.), The Miocene Land Mammals of Europe. Friedrich Pfeil, München. 557 Ginsburg, L. 2002. Les carnivores fossiles des sables de l’Orléanais. Annales de 558 Paléontologie 88:115–146. 559 Ginsburg, L. and C. Bulot. 1982. Les carnivores du Miocène de Bézian près de la 560 Romieu. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen 561 Serie B 85:53–76. 562 Ginsburg, L., and J. Morales. 1992. Contribution à la connaissance des Mustélidés 563 (Carnivora, Mammalia) du Miocène d’Europe Trochictis et Ischyrictis, genres affines 564 et genres nouveaux. Comptes Rendus de l’Académie des Sciences 315:111–116. 565 Gray, J. E. 1825. An outline of an attempt at the disposition of Mammalia into tribes 566 and families, with a list of the genera apparently appertaining to each tribe. Annals of 567 Philosophy 10:337–344. 568 Helbing, H. 1930. Zur Kenntnis der miocänen, “Mustela’’ zibethoides Blainville. 569 Eclogae Geologicae Helvetiae 23:637–644. 570 Hughes, S. S. 2012. Synthesis of Martes evolutionary history; pp. 3–22 in K.B. Aubry, 571 W. J. Zielinski, M. G. Raphael, G. Proulx, S. W. Buskirk (eds.), Biology and 572 Conservation of Martens, Sables, and Fishers. Cornell University Press, New York. 23 573 Jiangzuo, Q.-G., D. Gimranov, J.-Y. Liu, S.-Z. Liu, C.-Z. Jin, and J.-Y. Liu. 2021. A 574 new fossil marten from Jinyuan Cave, northeastern China reveals the origin of the 575 Holarctic marten group. Quaternary International 591:47–58. 576 Kretzoi, M. 1943. Kochictis centennii n. g. n. sp. az egeresi felsö oligocenböl. Földtani 577 Közlöny 73:10–17. 578 Larivière, S., and A. P. Jennings. 2009. Family Mustelidae (Weasels and Relatives); pp. 579 564–656 in D. E. Wilson and R. A. Mittermeier (eds.), Handbook of Mammals of the 580 World. 1. Carnivores. Lynx Editions, Barcelona, Spain. 581 Lartet, E. 1851. Notice Sur La Colline de Sansan, Suivie D’Une Recapitulation Des 582 Diverses Espèces D’Animaux Vertébrés Fossiles, Trouvés Soit À Sansan, Soit Dans 583 D’Autres Gisements Du Terrain Tertiaire Miocène Dans Le Bassin Souspyrénéen. J. 584 A. Portes, Auch, 60 pp. 585 Li, B., M. Wolsan, D. Wu, W. Zhang, Y. Xu, and Z. Zeng. 2014. Mitochondrial 586 genomes reveal the pattern and timing of marten (Martes), wolverine (Gulo), and 587 fisher (Pekania) diversification. Molecular Phylogenetics and Evolution 80:156–64. 588 Linnaeus, C. 1758. Systema Naturae Per Regna Tria Naturae, Secumdum Classes, 589 Ordines, Genera, Species, Cum Characteribus, Differentiis, Synonymis, Locis. 590 Laurentius Salvius, Stockholm, 823 pp. 591 Linnaeus, C. 1766. Systema naturae per regna tria naturae, secundum classes, ordinus, 592 genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio 593 duodecima, reformata. Laurentius Salvius, Stockholm. 532 pp. 594 Mayet, L. 1908. Études de Mammifères miocènes des sables de l’Orléanais et des faluns 595 de la Touraine. Annales de l'Université de Lyon 24:1–336. 596 Mein, P. 1958. Les mammifères de la faune sidérolithique de Vieux-Collonges. 597 Nouvelles Archives du Muséum d’Histoire naturelle de Lyon 5:1–122. 24 598 Meyer von H. 1842. Mittheilungen an Professor Bronn gerichtet. Neues Jahrbuch für 599 Geologie und Paläontologie 1842:583–589. 600 Montoya, P., J, Morales, and J. Abella. 2011. Musteloidea (Carnivora, Mammalia) 601 from the late Miocene of Venta del Moro (Valencia, Spain). Estudios Geológicos 602 67:193–206. 603 Morales, J., J. L. Cantalapiedra, A. Valenciano, D. Hontecillas, S. Fraile, B. A. García 604 Yelo, P. Montoya, and J. Abella. 2015. The fossil record of the Neogene Carnivore 605 Mammals from Spain. Palaeobiodiversity and Palaeoenvironments 95:373–386. 606 Morales, J., P. Peláez-Campomanes, P. Pérez, M. T. Alberdi, B. Azanza, M. Pickford, 607 M. Ríos, O. Sanisidro, G. Alcalde, J. L. Cantalapiedra, S. Fraile, B. García Yelo, A. 608 R Gómez Cano, V. Hernández-Ballarín, A. Oliver, E. Cantero, A. Valenciano, and P. 609 Montoya. 2018. Neogene Mammal sites from Molina de Aragón (Guadalajara, 610 Spain): correlation to other karstic sites of the Iberian Chain, and Geoheritage values. 611 Geoheritage 10:353–362. 612 Morlo, M. 1997. Die Raubtiere (Mammalia, Carnivora) aus dem Turolium von 613 Dorn-Dürkheim 1 (Rheinhessen). Teil 1: Mustelidae, Hyaenidae, Percrocutidae, 614 Felidae. Courier Forschungs-Institut Senckenberg 197:11–47. 615 Morlo, M., A. Le Maitre, K. Bastl, T. Engel, H. Lutz, B. Lischewsky, A. von Berg, 616 and D. Nagel. 2021. First record of the mustelid Trochictis (Carnivora, 617 Mammalia) from the early Late Miocene (MN 9/10) of Germany and a re- 618 appraisal of the genus Trochictis. Historical Biology 33:1183–1195. 619 Nagel, D., C. Stefen, and M. Morlo. 2009. The carnivoran community from the 620 Miocene of Sandelzhausen (Germany). Paläontologische Zeitschrift 83:151–174. 25 621 Peigné, S. 2012. Les Carnivora de Sansan; pp559–660 in: S. Peigné, and S. Sen (eds.), 622 Mammifères de Sansan. Mémoires du Muséum national d'Histoire naturelle. Paris: 623 Publications du Muséum, 203. 624 Peigné, S., M. J. Salesa, M. Antón, and J. Morales. 2006. New data on carnivores from 625 the Middle Miocene (Upper Aragonian, MN6) of Arroyo del Val area (Villafeliche, 626 Zaragoza Province, Spain). Estudios Geológicos 62:359–373. 627 Petter, G. 1963.Contribution à l’étude des Mustélidés des bassins néogènes du Vallès- 628 Penedès et de Calatayud-Teruel (Espagne orientale). Mémoires de la Société 629 géologique de France 43:1–44. 630 Petter, G. 1967. Mustélidés nouveaux du Vallésien de Catalogne. Annales de 631 Paléontologie 53:93–113. 632 Petter, G. 1976. Étude d’un nouvel ensemble de petits carnivores du Miocène 633 d’Espagne. Géologie Méditerranéenne 3:135–154. 634 Pinel, P. 1792. Recherches sur une nouvelle méthode de classification des quadrupèdes, 635 fondée sur la structure méchanique des parties osseuses qui servent à l’articulation de 636 la mâchoire inférieure. Actes de la Societé d’Histoire naturelle de Paris 1:50–60. 637 Pons-Moyà, J. 1990. Presencia de carnívoros turolienses en el Vallesiense terminal 638 (MN10) de Terrassa (Catalunya). Paleontologia i Evolució 23:199–203. 639 Roger, O. 1900. Wirbeltierreste aus dem Dinotheriensande. Naturwissenschaftlichen 640 Vereins für Schwaben und Neudorf in Augsburg 34:53–70. 641 Roth, C. and P. Mein. 1987. Martes collongensis n. sp. aus der miozänen Fundstelle 642 Vieux-Collonges (Dépt. Rhône, Frankreich). Mainzer geowissenschaftliche 643 Mitteilungen 16:157–164. 26 644 Samuels, J. X. and J. Cavin. 2013. The earliest known fisher (Mustelidae), a new 645 species from the Rattlesnake Formation of Oregon. Journal of Vertebrate 646 Paleontology 33: 448-454. 647 Samuels, J.X., K. E. Bredehoeft, and S. C. Wallace. 2018. A new species of Gulo from 648 the early Pliocene Gray fossil site (eastern United States); rethinking the evolution of 649 wolverines. PeerJ 6: p.e4648. 650 Schmidt-Kittler, N. 1981. Zur Stammesgeschichte der marderverwandten 651 Raubtiergruppen. Eclogae Geologicae Helvetiae 74:753–801. 652 Smith, J. B., and P. Dodson. 2003. A proposal for a standard terminology of anatomical 653 notation and orientation in fossil vertebrate dentitions. Journal of Vertebrate 654 Paleontology 23:1–12. 655 Stach, J. 1959. On some Mustelinae from the Pliocene bone breccia of Węże. Acta 656 Palaeontologica Polonica 4:101–117. 657 Sato, J. J., T. Hosoda, M. Wolsan, K. Tsuchiya, M. Yamamoto, and H. Suzuki. 2003. 658 Phylogenetic relationships and divergence times among mustelids (Mammalia: 659 Carnivora) based on nucleotide sequences of the nuclear interphotoreceptor retinoid 660 binding protein and mitochondrial cytochrome b genes. Zoological Science 20:243– 661 264. 662 Sesé, C. 1980. Mamíferos del Mioceno de Escobosa de Calatañazor (Soria). Ph.D. 663 dissertation, Universidad Complutense de Madrid, Madrid, 388 pp. 664 Valenciano, A. 2017: Mofetas, martas, tejones y rateles gigantes del Cerro de los 665 Batallones; pp. 322–336 in J. Morales (ed), La Colina de los Tigres Dientes de Sable. 666 Los Yacimientos Miocenos del Cerro de los Batallones (Torrejón de Velasco, 667 Comunidad de Madrid). Museo Arqueológico Regional, Madrid. 27 668 Valenciano, A., A. Pérez-Ramos, J. Abella, and J. Morales. 2020a. A new 669 hypercarnivorous mustelid (Mammalia, Carnivora, Mustelidae) from Batallones, late 670 Miocene (MN10), Torrejón de Velasco, Madrid, Spain. Geodiversitas 42:103–121. 671 Valenciano, A., J. Abella, D. M. Alba, J. M. Robles, M. A. Álvarez-Sierra, and J. 672 Morales. 2020b. New early Miocene material of Iberictis, the oldest member of the 673 wolverine lineage (Carnivora, Mustelidae, Guloninae). Journal of Mammalian 674 Evolution 27:73–93. 675 Van Dam, J. A. 2006. Geographic and temporal patterns in the late Neogene 12-3Ma 676 aridification of Europe: the use of small mammals as paleoprecipitation proxies. 677 Palaeogeography, Palaeoclimatology, Palaeoecology 238:190–218. 678 Van Dam, J. A., W. Krijgsman, H. Abels, M. A. Álvarez-Sierra, I. García-Paredes, P. 679 López-Guerrero, P. Peláez-Campomanes, and D. Ventra. 2014. Updated chronology 680 for the Middle to Late Miocene mammal sites of the Daroca Area (Calatayud- 681 Montalbán Basin, Spain). Geobios 47:325–334. 682 Villalta Comella J. F. de, and M. Crusafont Pairó. 1943. Los vertebrados del Mioceno 683 continental de la cuenca del Vallès-Penedès (provincia de Barcelona). I. Insectívoros. 684 II. Carnívoros. Boletín del Instituto Geológico y Minero de España 56:145–336. 685 Viret, J. 1933. Contribution à l’étude des Carnassiers miocènes de La Grive-Saint- 686 Alban (Isère). Travaux du Laboratoire de Géologie de la Faculté des Sciences de 687 Lyon 21:1–31. 688 Viret, J. 1951. Catalogue critique de la faune des Mammifères miocènes de La Grive 689 Saint-Alban (Isère). Première partie. Chiroptères, Carnivores, Édentés pholidotes. 690 Nouvelles Archives du Muséum d’Histoire naturelle de Lyon 3:1–104. 691 Wang, X., Z. J., Tseng, and G. T. Takeuchi. 2012. Zoogeography, Molecular 692 Divergence, and the Fossil Record-the Case of an Extinct Fisher, Pekania 28 693 palaeosinensis (Mustelidae, Mammalia), from the Late Miocene Baogeda Ula 694 Formation, Inner Mongolia. Vertebrata PalAsiatica 50:293–307. 695 Werdelin, L. 2005. The carnivores of Rudabánya; pp. 163–180 in: R. L. Bernor, L. 696 Kordos and L. Rook (eds.), Multidisciplinary Research at Rudabánya. 697 Palaeontographia Italica 90, Italy. 698 Werdelin, L., and S. Peigné. 2010. Carnivora; pp. 603–657 in L. Werdelin and W. 699 Sanders (eds.), Cenozoic Mammals of Africa. University of California Press, 700 Berkeley, California. 701 Zachos, J.C., Dickens, G.R., and Zeebe, R.E. 2008. An early Cenozoic perspective on 702 greenhouse warming and carbon-cycle dynamics. Nature 451:279–283. 703 Zdansky, O. 1924. Jungtertiäre carnivoren Chinas. Palaeontologica Sinica 2:1–149. 704 Submitted July 27, 2021; revisions received August 27, 2021; accepted Month DD, 705 YYYY 706 FIGURE CAPTIONS 707 FIGURE 1: Geographic location of fossil localities with Aragonictis araid gen. et sp. 708 nov. in the Iberian Peninsula. Abbreviations: 1, Escobosa de Calatañazor (Almazan 709 Basin); 2, Toril 3A (Calatayud-Daroca Basin); and 3, Andurriales (Calatayud-Daroca 710 Basin). Modified from Morales et al. (2018). [planned for 2/3 page width] 711 712 FIGURE 2: Mandibles and lower dentition of Aragonictis araid gen. et sp. nov. from 713 Andurriales, Toril 3A, and Escobosa de Calatañazor. A–C, MNCN 150.003 (holotype) 714 from Andurriales, left fragmentary hemimandible in A, buccal, B, lingual and C, 715 occlusal views; D–I, MNCN 150.004, right fragmentary hemimandible with m1 (D-F) 716 and p2-3 (G-I) from Andurriales in D, buccal, E, lingual, F, occlusal, G, buccal, H, 717 lingual, and I, occlusal views; J–L, MPZ 2020/474, left dp4 from Toril 3A in J, buccal, 718 K, lingual, and L, occlusal views; M–O, MNCN 26577, right fragmentary 29 719 hemimandible, screenshots of the 3D model of the CT scans from Escobosa de 720 Calatañazor in M, buccal, N, occlusal, and O, lingual views. [planned for full-page 721 width] 722 723 FIGURE 3: Maxillary and upper dentition of Aragonictis araid gen. et sp. nov. from 724 Escobosa de Calatañazor and Toril 3A. A MNCN 150.007, left fragmentary maxillary 725 from Escobosa de Calatañazor. Screenshots of the 3D model from CT scans in stereo 726 occlusal views; B, MPZ 2020/482, left fragmentary M1 from Toril 3A in occlusal view. 727 [planned for column width] 728 729 FIGURE 4: Bivariate plot of the length (L) and width (W) of the m1 of Neogene 730 marten-like mustelids and the extant Martes foina (type species of Martes). The linear 731 regression a’ (r2 =0.90) is defined by Miocene "Martes", Martes melibulla and Pekania 732 palaeosinensis, mostly characterized by their robust m1 talonid. The linear regression b’ 733 (r2 = 0.04) is defined by a sample of 19 specimens of M. foina. The linear regression c’ 734 (r2 = 0.85) is defined by relatively hypercarnivorous mustelids, characterized by slender 735 m1 and comprises Aragonictis araid, Circamustela spp., and Sinictis dolichognathus. 736 Values given in mm (Log). Abbreviations: r2, coefficient of determination; Log, 737 logarithm; 1, variability range of selected M. foina; 2, variability range of "Martes" 738 munki from numerous localities of Europe (MN4-MN8). The m1s pictures are showed 739 unscaled between them. Metric data from: Mayet (1908); Zdansky (1924); Villalta 740 Comella and Crusafont Pairó (1943); Dehm (1950); Viret (1951); Mein (1958); Petter 741 (1963,1967, 1976); Beaumont (1974); Ginsburg and Bulot (1982); Roth and Mein 742 (1987); Pons-Moyà (1990); Azanza et al. (1993); Ginsburg (2002); Peigné et al. (2006); 743 Nagel et al. (2009); Peigné (2012); Valenciano et al. (2020a) and this work. [planned for 744 full-page width] 30 745 746 FIGURE 5: Bivariate plot of the length (L) and width (W) of the M1 of Neogene 747 marten-like mustelids and the extant Martes foina (type species of Martes). Values 748 given in mm (Log). The ellipse marks the variability range of selected living M. foina 749 based on 20 specimens. Metric data from: Zdansky (1924); Dehm (1950); Viret (1933, 750 1951); Mein (1958); Petter (1976); Morlo (1997); Peigné (2012); Valenciano et al. 751 (2020a), and this work. [planned for full-page width] 752 753 FIGURE 6: Dentognathic remains of “Martes” cadeoti from Vieux-Collonges (middle 754 Miocene, MN5, France) and both Circamustela? laevidens and unknown Mustelidae 755 indeterminate from Wintershof-West (early Miocene, MN3, Germany). A–D, “Martes” 756 cadeoti from Vieux-Collonges. A–C, UCBL-FSL 65759, right m1 (holotype) in A, 757 buccal, B, lingual and C, occlusal views; D, UCBL-FSL 65758, left M1 (paratype) in 758 occlusal view; E–N, Circamustela? laevidens from Wintershof-West. E–G, SNSB- 759 BSPG 1937II 13306, right hemimandible with p4-m1 (holotype) in E, buccal, F, lingual 760 and G, occlusal views; H–J, SNSB-BSPG 1937II 13307, left hemimandible with p2, p4- 761 m1 in H, buccal, I, lingual and J, occlusal views; K, SNSB-BSPG 1937II 13902, right 762 hemimandible with p2-4, and a fragmented m1; L. SNSB-BSPG 1937II 13372, right 763 maxillary with P4-M1 in occlusal view; M. SNSB-BSPG 1937II 13722, left P4 in 764 occlusal view; N. SNSB-BSPG 1937II 13724, left M1 in occlusal view; O. Unknown 765 Mustelidae from Wintershof-West. SNSB-BSPG 1937II 13723, right M1 in occlusal 766 view. [planned for full-page width] 31