This is the accepted version of the journal article: Bartolini-Lucenti, Saverio; Madurell-Malapeira, Joan; Rook, Lorenzo. «The carnivorans from Cava Monticino (Faenza, Italy; Messinian) revisited». Historical Biology, (February 2022). DOI 10.1080/08912963.2022.2042806 This version is available at https://ddd.uab.cat/record/257658 under the terms of the license 1 The carnivorans from Cava Monticino (Faenza, Italy; Messinian) revisited 2 Saverio Bartolini-Lucenti1*, Joan Madurell-Malapeira2, Lorenzo Rook1 3 4 1 Università degli Studi di Firenze, Dipartimento di Scienze della Terra, Via G. La Pira 4, 50121 Firenze (FI) – 5 saverio.bartolinilucenti@unifi.it, lorenzo.rook@unifi.it 6 2 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, C\ de les Columnes sn, Campus de la UAB, 7 08193 Cerdanyola del Vallès, Barcelona, Spain – joan.madurell@icp.cat 8 9 Corresponding Author: saverio.bartolinilucenti@unifi.it 10 11 Abstract 12 Among the vertebrates found at Cava Monticino, carnivorans are by far the most abundant of all the 13 large mammals. Five different taxa were recovered: one felid, two hyaenids, one canid, and one 14 mustelid. The small-sized felid remains can be attributed to Felis christoli and seems to represent 15 one of the earliest records of a true member of the genus Felis in Western Europe. Hyaenids at Cava 16 Monticino are represented by the large wolf-sized and cursorial Lycyaena cf. chaeretis, and by the 17 peculiar small Plioviverrops faventinus, the most abundant taxon of all. The latter is one of the most 18 derived species of the genus and the last to appear in the fossil record of these mongoose-like 19 hyaenids. The medium-sized canid recorded at Cava Monticino, Eucyon monticinensis, represent 20 one of the oldest, certain record in the Old World of the genus Eucyon. It was a mesocarnivorous 21 species that preyed on small vertebrates (abundantly recorded in the area of Cava Monticino during 22 the Late Miocene). Lastly, mustelids are represented by the large relative of the extant honey 23 badger, Mellivora benfieldi, whose record at Cava Monticino represents the northernmost record of 24 the species and, presently, the only record of the genus outside of Africa. 25 26 Keywords: 27 Felidae, Hyaenidae, Canidae, Mustelidae, Late Miocene, Messinian. 28 29 Introduction 30 Terrestrial carnivores are an important ecological component in a wide range of biomes, and overall 31 contribute to characterizing faunal assemblage ecology, composition, and structure. In respect to the 32 general structure of other well-known Late Miocene faunas (such as Pikermi and Samos in Greece 33 or Batallones and Venta del Moro in Spain) where the number of recorded species is similar to the 34 high taxonomic diversity seen in modern African or Asian faunas, the carnivore guild documented 35 at Cava del Monticino assemblage is composed by only five taxa (Rook et al. 1991): one felid (the 36 small cat referred to Felis ex gr. attica-christoli; two hyaenids (the small mongoose-like 37 Plioviverrops faventinus, and the large-sized meat-eater hyaenid, referred to Thalassictis 38 (Lycyaena) gr. macrostoma-chaeretis); one canid (the primitive dog Eucyon monticinensis, and one 39 mustelid (the honey badger Mellivora benfieldi). Despite its reduced composition, the carnivorans 40 from Cava Monticino represent one of the best and most abundant association of carnivorans dated 41 to the latest Miocene of Italy and Western Europe, in general. Here we provide an updated revision 42 of these carnivorans, with particular attention to the most problematic ones (the small felid and the 43 large hyaenid) and focusing on the significance of Cava Monticino guild in terms of paleoecology, 44 biochronology and zoogeography. 45 46 Materials & Methods 47 The present study is carried out studying the material of carnivorans housed at the “Museo Civico di 48 Scienze Naturali Malmerendi” of Faenza (Emilia Romagna, Italy) and some isolated teeth and 49 fragmented specimens from the Earth Science Department of the University of Florence. For the 50 Miocene hyaenid discussion, we followed the ecomorphological interpretation by Werdelin and 51 Solounias (1991) and the taxonomic revision by Turner et al. (2008). Metric and morphological 52 comparison was taken by literature (inter alios De Beaumont 1961; Ficcarelli and Torre 1975; 53 Morales and Aguirre 1976; de Bonis and Koufos 1991; Werdelin et al. 1994; Werdelin 1999; Salesa 54 et al. 2012, 2019; Robles et al. 2013; Ozkurt et al. 2015; Koufos 2021). Measurements were taken 55 to the nearest 0.1 mm with a digital caliper, following Driesch (1976) and reported in the 56 Supplementary Material. 57 58 Systematic Paleontology 59 Order Carnivora Bowdich, 1821 60 Family Felidae Fischer, 1817 61 Sub-Family Felinae Fischer, ,1817 62 Genus Felis Linnaeus, 1758 63 64 Felis christoli Gervais in Gervais, 1848-1852 65 Figure 1A-B, I 66 Material – Mandible. MSF 90 (BRS 27/3), right hemimandible fragment with partial p4 and m1. 67 Postcranial elements. MSF 91 (BRS 27/12), third left metatarsus; MSF 467 (BRS 27/sn), four left 68 metatarsus. 69 70 Remarks – The generic attribution of the species F. christoli has been questioned, relating it to 71 extant forms, like the serval, Leptailurus serval (see Gervais 1859), to lynxes (see Morales and 72 Aguirre 1976), to Pseudaelurus or even Metailurus (see Ficcarelli and Torre 1975). Other authors 73 left the question open (Werdelin et al. 2010), considering the scantiness of its fossil record. 74 Nevertheless, the comparison of the mandible (e.g., the angle between the ramus and the corpus; 75 reduction of the angular process; height of the condyloid process) and of the m1 (e.g., absence of 76 the talonid; proportion between paraconid and protoconid) with those of the members of the above- 77 mentioned fossil and extant genera does not show any affinity, if not with Felis sensu stricto. The 78 phylogenetic relationships and evolutionary history of Miocene felids have been a harshly debated 79 topic in the last century as a consequence of their scarce record especially in the Early-Middle 80 Miocene (see Werdelin et al. 2010 for discussion). We report a resuming map of the occurrences of 81 small felids from the MN10 onward (Figure 2). Some authors, in spite of the initial attribution of 82 Miocene small felid to the genus Felis (sensu lato), have preferred including these forms into 83 different genera (e.g., Salesa et al. 2012, 2019). 84 Compared to the old forms attributed to the genera Pseudaelurus, e.g., Pseudaelurus romieviensis 85 (type locality: La Romieu, France, MN4), P. quadridentatus (type locality: Sansan, France, MN6) 86 and P. turnauensis (type locality: Göriach, Austria, MN5), MSF 90 lacks the developed talonid on 87 the m1, evident In these primitive forms (De Beaumont 1961; Robles et al. 2013) (Figure 1). This 88 same relevant difference strongly suggests the distinction between the specimen from Cava 89 Monticino and the Spanish MN10 Leptofelis vallesiensis (see Salesa et al. 2019) in which the 90 talonid is as marked as in small and primitive Pseudoaelurus s.l. Moreover, L. vallesiensis shows a 91 short m1 paraconid compared to the protoconid. Several morphologies of the type and hypodigm 92 material of the MN12 Felis attica from Pikermi and Samos allow separating the fossils from Cava 93 Monticino from this small felid (de Beaumont 1961; Koufos 2011). For instance, in F. attica, the 94 corpus of the mandible is shallower with an inclined ramus, proportionally higher coronoid process 95 but a shorter height between the angular process and the condyloid process. The angular process is 96 smaller and more slender in MSF 90 (Figure 1A-B). Even in dental morphologies, there are 97 differences: F. attica shows a shorter blade of the paraconid compared to the protoconid, unlike 98 MSF 90, in which the two cuspids are equally developed (Figure 1). Moreover, MSF 90 shows a 99 very small, but evident, metaconid, whereas F. attica does not. Like some specimens of F. attica, 100 MSF 90 does not have a talonid, nor distal enlargement or cingulid. For what is visible, the distal 101 portion of the p4 differs from that of F. attica for the smaller distal accessory cuspulid and less 102 sharp cingulid; moreover, in occlusal view, the distal portion of the p4 of MSF 90 is more enlarged 103 compared to that of F. attica (Figure 1). The overall morphology of the lower m1, with subequal 104 paraconid and protoconid, recalls the morphologies of the MN13-14 Felis christoli recovered from 105 the marine sands of Montpellier and described from Venta del Moro (Gervais 1859; Ficcarelli and 106 Torre 1975; Morales and Aguirre 1976). Other similarities are evident like the position of the 107 mental foramina, the general shape of the m1, with a bulging distobasal portion, instead of a 108 talonid; a small pointy metaconid; and the sharp buccal cingulids on the p4 and the m1. The type 109 mandible is rather shallow and possesses a considerably stout distal accessory cuspulid on the p4 110 and a visible gap between paraconid and protoconid on the m1, although such gap is not present in 111 MP338 (one of the two specimens housed in the Museum of Basel). More importantly, the available 112 specimens of F. christoli do not have any trace of a metaconid, unlike the greatly reduced one of 113 MSF 90. Altogether, the set of features suggests an attribution to a form close to the French and 114 Spanish late Miocene-Early Pliocene F. christoli. This interpretation is supported also by the similar 115 size between the specimens of F. christoli and those of Cava Monticino, intermediate between large 116 lynxes and smaller feral cats (Figure 3). According to the above-mentioned morphology and overall 117 proportions, we include the Cava Monticino specimens in Felis christoli. 118 119 Family Hyaenidae Gray, 1821 120 Sub-Family Ictitheriinae Dietrich, 1927 121 Genus Plioviverrops Kretzoi, 1938 122 123 Plioviverrops faventinus Torre, 1989 124 Figure 4A-B, Figure 5A-B 125 126 Material – MSF 62, fossiliferous block with several specimens. 127 Cranium. MSF 408 (BRS 5/33), palate with left and right P4; MSF 430 (BRS 19/5), left maxillary fragment 128 P3-P4. 129 Upper teeth. MSF 423_1 (BRS 5/294), right C; MSF 411 (BRS 5/61), right C; MSF 417 (BRS 5/278), right 130 P2; MSF 423_1 (BRS 5/279), left P2; MSF 424_1 (BRS 5/279), left P2; MSF 424_2 (BRS 5/279), left P2; 131 MSF 418 (BRS 5/300), right P3; MSF 425_1 (BRS 5/64), right P3; MSF 425_2 (BRS 5/65), left P3; MSF 132 425_3 (BRS 5/301), right P3; MSF 425_4 (BRS 5/281), left P3; MSF 426_1 (BRS 5/63), left P3; MSF 415 133 (BRS 5/165), right P4; MSF 424_2 (BRS 5/297), right P4; MSF 424_3 (BRS 5/162), left P4; MSF 413 (BRS 134 5/313), left M1; MSF 428_1 (BRS 5/310), right M1; MSF 428_2 (BRS 5/56), right M1; MSF 428_3 (BRS 135 5/sn), right M1; MSF 414_1 (BRS 5/57), right M2; MSF 414_2 (BRS 5/312), right M2. 136 Mandible. MSF 92 (BRS 5/34), right hemimandible with p3-m2; MSF 92.1 (BRS 5/34), left hemimandible 137 fragment with p4-m2; MSF 93 (BRS 5/30), left hemimandible with c and p3; MSF 94 (BRS 19/6), right 138 edentulous hemimandible; MSF 407 (BRS 5/158), right hemimandible fragment with p3-m2. 139 Lower teeth. MSF 412 (BRS 5/sn), left c; MSF 423_1 (BRS 5/16), left c; MSF 423_2 (BRS 5/293), right c; 140 MSF 423_3 (BRS 5/62), right c; MSF 423_4 (BRS 5/279), right c; MSF 409 (BRS 5/167), right p3; MSF 141 410 (BRS 5/168), right p4; MSF 416 (BRS 5/54), left m1; MSF 417_1 (BRS 5/169), right m1; MSF427_1 142 (BRS 5/314), left m1;MSF 427_2 (BRS 5/sn), right m1; MSF 427_3 (BRS 5/67), right m1; MSF 427_4 143 (BRS 5/68), right m1; MSF 429_1 (BRS 5/sn), left m2; MSF 429_2 (BRS 5/sn), right m2; MSF 429_3 (BRS 144 5/sn), left m2; MSF 429_4 (BRS 5/170), right m2; MSF 429_5 (BRS 5/286), right m2. 145 Remarks – Remains of a small-sized hyaenid are among the most abundant of all mammals 146 recovered at Cava Monticino. This important record features also some impressive fossils (Figure 147 1). The species was described extensively by late Danilo Torre, Professor of the University of 148 Florence, in 1989 who was the first to acknowledge the peculiarity of the teeth morphologies of the 149 Cava Monticino small hyaenid (Torre 1989). The cranium morphology, although crushed and 150 deformed in the specimens from Cava Monticino, is somewhat in between that of herpestids and 151 that of a fox, with a longer neurocranium compared to the snout. It has some features that recall 152 modern herpestids, for instance, the posterior portion of the mandible in lateral view or the full 153 dentition with four premolars and two upper and lower molars. Cava Monticino Plioviverrops 154 displays crushing-puncturing cusps on its teeth, especially on the molars, like other species of the 155 genus across Europe. For instance, compared to Plioviverrops guerini and Plioviverrops orbignyi 156 the P4 has a mesiodistally shorter metastyle and, in general, a proportionally larger surface of the 157 upper P4 (for the higher cingulum; the lingually expanded paracone and protocone); the 158 morphology of the M1 is similar in three species but P. faventinus has a slender M2, compared to 159 the squared one of P. guerini or of P. orbignyi (Alcalá 1994; de Bonis and Koufos 1991; Koufos 160 2011). Compared to both P. guerini and P. orbignyi, the lower premolars of P. faventinus have 161 more developed accessory cuspulids, with larger distal portions, which in the case of the p3 and p4 162 have the shape of talonid (Torre 1989). Torre (1989) regarded P. orbignyi and P. guerini as closely- 163 related species, zoogeographic variants of single species dwelling Southern Europe, and P. 164 faventinus as derived from P. orbignyi. In terms of size, P. faventinus is closer to P. guerini, as P. 165 orbignyi is overall smaller (Figure 6). Nevertheless, in teeth proportions, the similarity of P. 166 faventinus lies closer to P. orbignyi as P. guerini has slender carnassials and upper molars and a 167 proportionally larger m2. These similarities and differences, rather than a true phylogenetic affinity 168 might be a hint of a similar ecological adaptations (as seen in other carnivorans, Bartolini-Lucenti 169 and Rook 2020). 170 Although Plioviverrops is among the earliest hyaenids to appear in the fossil record, as its earliest 171 occurrences are in the MN4-5 (ca. 16.5 Ma) (Turner et al. 2008), Plioviverrops faventinus is the 172 youngest species of this mongoose-like genus to appear in the fossil record (Figure 7). The earliest 173 records of Plioviverrops are scanty and not well characterized (Turner et al. 2008). Plioviverrops 174 orbignyi from numerous localities of Greece dated between 9.7-7.0 Ma (Koufos 2011); and P. 175 guerini from Spanish and Greece MN11-12 (8.7-7.0 Ma) sites (Alcalá 1994; Koufos 2011) (Figure 176 7). The extensive record of Cava Monticino makes Plioviverrops faventinus one of the best-known 177 species of the genus. As visible from Figure 6, P. faventinus is overall larger compared to P. 178 orbignyi and it is more similar to P. guerini in this respect. Whereas P. orbignyi weighted around 3- 179 4 kg, both P. guerini and P. faventinus had a probable body mass of ca. 7 kg. Moreover P. 180 faventinus is one of the latest species of ecomorph Group 2 to survive the Miocene-Pliocene 181 transition, together with the P. cf. orbignyi from Çalta-1 (Sen and Saraç 2018). Fossils of P. 182 faventinus have been also recovered from the Spanish site of La Gloria 4 (ca. 4.1 Ma). 183 184 Sub-family Hyaeninae Mivart, 1882 185 Genus Lycyaena Hensel, 1863 186 187 Lycyaena cf. chaeretis (Gaudry, 1861) 188 Figure 4C-D, Figure 5C-F 189 190 Material – Cranium. MSF 84 (BRS 27/17), cranium with right P3-M1. 191 Upper teeth. MSF 385 (BRS 27/19), a third upper left incisor; MSF 383 (BRS 8/9), left P3; MSF 401 (BRS 192 27/18), left M1. 193 Mandible. MSF 384 (BRS 27/1), juvenile left mandible. 194 Lower teeth. MSF 83 (BRS 25/20), upper left canine; MSF 86 (BRS 27/2), right p4; MSF 87, right m1 (BRS 195 27/28). 196 Postcranial elements. MSF 393, second left metacarpal (BRS 27/35); MSF 394 (BRS 27/36), third right 197 metacarpals; BRS 27/39, third right metacarpals; MSF 395, fifth left metacarpals (BRS 27/36); MSF 396, 198 fifth left metacarpals (BRS 27/37); MSF 64 (BRS 27/31), the proximal part of a left radius; MSF 526, right 199 fragment of pelvis; MSF 404, distal epiphysis of a juvenile right femur; MSF 403 (BRS 27/32), distal 200 fragment of a right tibia; MSF 389 (BRS 27/30), proximal fragment of a right tibia; BRS27/48 (BRS 27/48), 201 a juvenile left tibia; MSF 388 (BRS 27/29), right astragalus and lateral cuneiform; MSF 391(BRS 27/33), left 202 calcaneum; MSF 392 (BRS 27/34), right calcaneum; MSF 398, third metatarsal; MSF 399, first left phalanx; 203 BRS 27/9 (BRS 27/41), first right phalanx; MSF 400 (BRS 27/42), first left phalanges; BRS 27/43, second 204 left phalanges; BRS 27/44, second left phalanges. 205 206 Remarks – The morphologies possessed by the large-sized hyaenid from Cava Monticino are a 207 mixture of derived and primitive features for late Miocene Hyaenidae (Figures 4-5): large 208 mesiolingual cuspule on the P3; large paracone on the P4; presence of M1 but absence of the M2; M1 209 triangular-shaped; the presence of an individualized well developed distolingual cuspulid on the p4; 210 rectangular distal part of the p4; m1 talonid modestly developed, with one large hypoconid and a 211 ridge-like, lingual entoconid; small m1 metaconid (Figure 4-5). A peculiarity of the large hyaenid of 212 Cava Monticino is the prominently developed mesial cuspulid on the P3. The set of morphologies 213 possessed by the specimens of Cava Monticino contrasts with the features of Thalassictis s.l., e.g., 214 the dorsoventrally shallow cranium, unlike Thalassictis robusta from Kishinev (see Semenov 2008), 215 Thalassictis montadai from Hostalets de Pierola; the presence of the M2 (as in T. robusta from 216 Kishinev); the absence of a lingual cuspulid on the p4; the larger m1 talonid with developed 217 hypoconid and entoconid. The presence of an m2, is not directly testable on the sample of Cava 218 Monticino, as no m2 nor any mandible of a large hyaenid has been found. The only mandible 219 preserved at Cava Monticino is that of a very young individual with a dp4, but the x-rays image on 220 the mandible has revealed only the roots of the dp4 and the m1, and no m2. These features suggest 221 discarding an attribution to Thalassictis. Despite the similarity in size, especially with Hyaenictis 222 hendey (Figure 8), an attribution to Hyaenictis seems improbable as one of the major diagnostic 223 features of the genus according to Werdelin et al. (1994), i.e., the presence of the m2, is absent in 224 Cava Monticino’s hyaenid. Compared to Hyaenictitherium wongii, the sample from Cava Monticino 225 is larger (Figure 8) and possesses several features contrasting with the specimens attributed to the 226 former species, e.g., the absence of the M2, the shortened m1, the reduced talonid cuspulids. MSF 84 227 has a shortened cranium, and especially the muzzle, and contrasts with that of some Lycyaena spp., 228 in which is generally more slender and elongated e.g., to Lycyaena macrostoma from Dhok Pathan 229 (Pilgrim 1932) and Lycyaena dubia from Hayranlı-Sivas (Ozkurt et al. 2015) and Lok. 49 (Zdansky 230 1924), but it resembles that of Lycyaena chaeretis from Samos (unknown locality; Koufos 2021). 231 Another similarity to the sample from Samos is the shape of the M1, although differences from this 232 specimen and from the type (recovered from Pikermi) should be highlighted. The hyaenid from Cava 233 Monticino has a large mesiolingual cuspule on the P3; lowered p4 protoconid contrasting with the 234 slender one of L. chaeretis; the p4 distolingual cuspulid is individualized and laterally positioned 235 compared to the distal cingulid; the m1 metaconid is more reduced, and the talonid only presents a 236 large centralized hypoconid and a cristid-like cuspulid on the lingual side. In size, the taxon from 237 Cava Monticino is really large, close to the largest known species of the genus L. crusafonti from 238 Bled Douarah (Tunisia) and to Lycyaena cf. chaeretis from Cerro de la Garita (Alcalá 1994), see 239 Figure 8. For this mixture of similarities and the relevant differences, we prefer to ascribe the taxon 240 from Cava Monticino to Lycyaena cf. chaeretis, conscious that more research needs to be carried out. 241 The genus Lycyaena has a Eurasian and North African distribution (Figure 9): it is known since the 242 MN 9 (a period comprised between 11 and 9.7 Ma) when the species L. crusafonti is recovered 243 from the locality 17 of Bled Douarah (Tunisia). Further occurrences are those of the MN 12 (ca 244 7.75-7.0 Ma): L. macrostoma in the Siwalik basin (India-Pakistan); L. dubia in several localities of 245 China; and L. chaeretis in the famous Greek Miocene sites of Pikermi and Samos. The occurrence 246 at Cava Monticino of this large-sized cursorial hunter is relevant for several reasons. Firstly, this 247 record represents the last occurrence of this genus in the Old World, separated from other 248 congeneric taxa by nearly 1.5 Myr. Furthermore, this of Cava Monticino is one of the largest 249 Lycyaena of the fossil record, even larger than the earliest taxon L. crusafonti (Figure 8). 250 251 Family Canidae Fischer, 1817 252 Subfamily Caninae Fischer, 1817 253 Tribe Canini Fischer, 1817 254 Genus Eucyon Tedford and Qiu, 1996 255 256 Eucyon monticinensis (Rook, 1992) 257 Figure 1C-F, K-L 258 259 Material – Cranium. MSF 466, cranium fragment. 260 Upper teeth. BRS 27/8, right upper canine; BRS 27/25, right upper canine; BRS 27/26, right upper canine; 261 BRS 27/51, right upper canine; BRS 24/12, right M1. 262 Mandible. MSF 97 (BRS 27/4), right hemimandible with p2 and m1; MSF 456 (BRS 27/47), a juvenile left 263 hemimandible with p2-p4 in eruption. 264 Lower teeth. BRS 27/27, three incisors; BRS 27/7, left p2; BRS 27/22, right p2; BRS 27/6, right m1; BRS 265 27/21, left m1; BRS 27/5; right m2; BRS 27/24; right m2. 266 Postcranial elements. MSF 100 (BRS 27/16), left humerus; MSF 98 (BRS 27/15), right radius; MSF 99 267 (BRS 27/14), left ulna; BRS 27/46, right fifth metacarpal; MSF 101 (BRS 27/13), right tibia; BRS27/45, left 268 calcaneum; BRS27/10, right astragalus. 269 270 Remarks – The medium-sized canid from Monticino has been described in deep by Rook (1992) as 271 Eucyon monticinensis. Its record at Cava Monticino is fairly abundant compared to the other taxa, 272 with cranial and postcranial elements of at least 5 individuals. The type, the right mandible MSF 97 273 with p2 and m1 (Figure 1E-F, L), shows all the peculiar features that support distinction of E. 274 monticinensis from other similar sized and coeval species, like the proportionally stout and deep 275 corpus, m1with developed trenchant portion but also developed talonid one (crushing surface). The 276 genus Eucyon is a rather diverse genus of canids that arrived in the Old World between at the end of 277 the Miocene (ca. 7 Ma) and disappeared during the Early Pleistocene (Figure 10). The occurrence 278 of E. monticinensis is really important in the Eurasian record as it is the second earliest canid to 279 reach Western Europe, after Eucyon cipio (Rook 2009) (Figure 10). Other occurrences of E. 280 monticinensis are those of Verduno (Alba, Piedmont) and of the upper cycle of Casino Basin 281 (Siena, Tuscany) (Figure 10). Surely, this species represents nowadays the only species of Eucyon 282 yet discovered in Italy. Rook (1992) recognized an affinity between E. monticinensis and the 283 sample of the medium-sized canid from Venta del Moro (a coeval site to Cava Monticino). The 284 latter has been recently revised and ascribed to the species Eucyon debonisi (see Montoya et al. 285 2009). The two sample differ for a slight difference in size, as E. debonisi is overall smaller (Figure 286 11). There are also some dentognathic features that distinguish the material from Venta del Moro 287 from that of Cava Monticino, buccolingual margin of the M1 (sharp in E. debonisi and rounded in 288 E. monticinensis), the robustness of the corpus (stronger in E. monticinensis), shape of the m1 289 (more slender and buccolingually compressed in E. debonisi, with a mesiodistally short paraconid 290 and an elongated talonid). Such features are evident although the scarcity of adequately abundant 291 material to assess the entity of intraspecific variability of those two forms leaves the question of 292 their real distinction open. 293 294 Family Mustelidae Swainson, 1835 295 Sub-Family Mellivorinae Gray, 1865 296 Genus Mellivora Storr, 1780 297 298 Mellivora benfieldi Hendey, 1978 299 Figure 1C, J 300 301 Material – Mandible. MSF 88 (BRS 25/19), left hemimandible with p3-m1. 302 Remarks 303 The extant honey-badger, Mellivora capensis, is the only living representative of Mellivorinae 304 although the fossil record of the subfamily is much more abundant (Valenciano and Govender 305 2020). The earliest occurrences of mellivorines are at MN7-8 (ca 12 Ma) from the Ngorora 306 formation in Kenya, and MN12 (ca. 7 Ma) from Toros Menalla 192 (Chad) and Lothagam (Kenya). 307 The genus Mellivora seems to appear around 6.0 Ma, and the species M. benfieldi it is indeed the 308 earliest species of the genus. It was described for the first time from the latest Miocene-earliest 309 Pliocene site of Langebaanweg (South Africa; Hendey 1978) (Figure 12). During the Pliocene, the 310 extant species M. capensis arose in Africa whereas in Eurasia and North America appeared the 311 large-sized genus Eomellivora (Valenciano and Govender 2020). The sample of Cava Monticino is 312 composed of a single left mandible with p3-m1. As Rook et al. (1991) noted the specimen is really 313 similar to the original material of M. benfieldi from Langebaanweg if not for minor dental features. 314 This interpretation was followed by a recent restudy by Valenciano and Govender (2020). The 315 record from in the Italian Messinian locality is relevant as it testifies to the northern-most 316 occurrence of this species in the Old-World and the first occurrence on the genus out of the African 317 continent (Figure 12). The only other occurrence of the species is the Late Miocene site of Middle 318 Awash. 319 320 Concluding remarks 321 The carnivoran assemblage recovered from Cava Monticino is, although limited in species richness, 322 relevant in terms of the peculiar composition and different ecology. Of the carnivorans there 323 recorded, the top predator was surely the large-sized Lycyaena cf. chaeretis. Although size 324 estimations for the other species of the genus are around 50 kg, the individuals of Cava Monticino 325 weighted more than 60 kg. Considering its morphological features (Rook et al. 1991 and the present 326 study) and enamel structure (Ferretti 2007), this hyaenid was interpreted as a hypercarnivorous 327 species referred to the ecomorph group 4 by Werdelin and Solounias (1991). Like others of this 328 ecomorph group, L. cf. chaeretis from Cava Monticino was a cursorial wolf-like, meat- and bone- 329 eater capable of hunting down large ungulates, like for instance hipparions, probably after tiring 330 them out with long runs, like modern canids do (the African hunting dog or wolves). Lycyaena cf. 331 chaeretis is also the only large-sized carnivorans yet recovered from Cava Monticino. The other 332 carnivorans are all small- or medium-sized. The felid Felis christoli was more similar to a modern 333 small lynx than to feline cats, and, therefore a similar hunting strategy to that of modern Lynx can 334 be hypothesized. The dental and mandibular morphology of Plioviverrops faventinus are closely 335 resembling those of extant Herpestidae, as identified by Werdelin and Solounias (1991) who 336 classifies the species of this genus in the ecomorph group 2 (mongoose-like Hyaenidae). Ferretti 337 (2007) remarked the omnivorous nature of Plioviverrops’s diet pointing out macroscopic and 338 microscopic features like the crushing-puncturing dentition and the outlines of the Hunter-Schreger 339 bands. Studying mandibular and dental proportions, Bartolini Lucenti and Rook (2020) found that 340 E. monticinensis probably was able to hunt larger prey compared to other Mio-Pliocene eucyons, up 341 to ca. 5 kg in size. The great variety of small mammals found at Cava Monticino might give us a 342 hint of the diet of this canid: from mice, dormice and hamsters to lagomorphs (like Prolagus or 343 Trischizolagus), and probably not distasting some herpetofaunal species as well (Villa et al. 2021). 344 E. monticinensis represents an important element of the fauna of Cava Monticino and also for other 345 European Messinian localities. This medium-sized canid probably occupied an ecological niche 346 intermediate between the small-sized Plioviverrops faventinus and the large-sized Lycyaena cf. 347 chaeretis, filling the empty place of medium-sized hyenas, as none of them have been discovered in 348 the fossil record of Cava Monticino. Ecologically speaking, M. benfieldi was probably very similar 349 to the extant M. capensis as few dental differences can be pointed out between the two. M. benfieldi 350 could have been a stout omnivorous carnivorans preying on small vertebrates like amphibians, 351 reptiles, birds, and mammals but also insects. Like the extant species it could also eat bulbs, berries, 352 and fruits. The peculiar affection of extant M. capensis for honey and bee larvae cannot be proved 353 in the fossil taxon but the possibility cannot be ruled out either. 354 355 Acknowledgments 356 We are deeply indebted to Antonio (“Tonino”) Benericetti for his passion and heartful dedication to 357 Cava Monticino paleontology, to Marco Sami and Enzo Bagnaresi (Museo di Storia Naturale di 358 Faenza) for access to collections under their care, and to Piero Lucci and Massimo Ercolani 359 (Federazione Speleologica Regionale dell’Emilia-Romagna) for the encouragement in starting a 360 revision of the Cava Monticino fauna. We thank Prof. J. Morales for his contributions in the 361 advance of Vertebrate Paleontology. This contribution is framed within a wider project on late 362 Neogene vertebrate evolution at the University of Florence (coordinator LR). 363 The present manuscript has been conceived and prepared during a Visiting Professor fellowships at 364 the Earth Sciences Department of the University of Florence awarded to JMM in May-June 2021. 365 This work has been also funded by the Agencia Estatal de Investigación-European Regional 366 Development Fund of the European Union (CGL2017-82654-P, AEI/FEDER-UE) and the 367 Generalitat de Catalunya (CERCA Program). J.M.-M. is member of the consolidated research group 368 2017 SGR 116 (AGAUR, Generalitat de Catalunya). 369 370 Disclosure statement 371 No potential conflict of interest was reported by the authors. 372 373 Funding 374 This paper has been supported by the University of Florence “Fondi di Ateneo” and “Fondi 375 internazionalizzazione” to LR. 376 377 ORCID 378 Saverio Bartolini-Lucenti http://orcid.org/0000-0003-1280-5378 379 Joan Madurell-Malapeira http://orcid.org/ 0000-0003-4639-9451 380 Lorenzo Rook http://orcid.org/0000-0001-8923-5428 381 382 References 383 Alcalà L. 1994. Macromamiferos neógenos de la fosa de Afambra-Teruel. Madrid: Instituto de 384 Estudios Turolenses: CSIC, Museo Nacional de Ciencias Naturales. 385 386 Bartolini Lucenti S, Rook L. 2020. "Canis" ferox Revisited: Diet Ecomorphology of Some Long 387 Gone (Late Miocene and Pliocene) Fossil Dogs. J Mammal Evol. 28(2):285–306. 388 389 De Beaumont G. 1961. 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New Fossils of Mellivora Benfieldi (Mammalia, Carnivora, 459 Mustelidae) from Langebaanweg, ‘E’ Quarry (South Africa, Early Pliocene): Re-Evaluation 460 of the African Neogene Mellivorines. J Vert Paleontol. 40: e1817754. 461 462 Valenciano A, Morales J, Govender R. 2021 Eucyon khoikhoi sp. nov.(Carnivora: Canidae) from 463 Langebaanweg ‘E’Quarry (early Pliocene, South Africa): the most complete African canini 464 from the Mio-Pliocene. Zool J Linn Soc. https://doi.org/10.1093/zoolinnean/zlab022 465 466 Villa A, Carnevale G, Pavia M, Rook L, Sami M, Szyndlar Z, Delfino M. 2021. An overview on the 467 late Miocene vertebrates from the fissure fillings of Monticino Quarry (Brisighella, Italy), 468 with new data on non-mammalian taxa. Riv It Paleont Strat. 127:297–354. 469 470 Werdelin L. 1999. Studies of fossil hyaenas: affinities of Lycyaenops rhomboideae Kretzoi from 471 Pestlörinc, Hungary. Zool J Linn Soc. 126:307-17. 472 473 Werdelin L, Solounias N. 1991. The Hyaenidae: taxonomy, systematics and evolution. Fossils and 474 strata. 30:1–104. 475 476 Werdelin L, Turner A, Solounias N. 1994. Studies of fossil hyaenids: the genera Hyaenictis Gaudry 477 and Chasmaporthetes Hay, with a reconsideration of the Hyaenidae of Langebaanweg, South 478 Africa. Zool J Linn Soc. 111:197–217. 479 480 Werdelin L, Yamaguchi N, Johnson WE, O’Brien SJ. 2010. Phylogeny and evolution of cats 481 (Felidae). In Macdonald DW, Loveridge AJ, editors. Biology and conservation of wild felids. 482 Oxford: Oxford University Press; p. 59-82. 483 484 Zdansky O. 1924. Jungtertiäre Carnivoren Chinas. Palaeontol Sin. 2: 1–149. 485 486 Figure Caption 487 488 Figure 1 Carnivoran materials from Cava Monticino (Brisighella, RA, Italy). Felis christoli: A-B, I - MSF 90 (BRS 489 27/3), right hemimandible with p4-m1 in lateral (A), occlusal (B) and detailed occlusal photographic and 3D model 490 views (I). Eucyon monticinensis: C-F, K-L – BRS 24/12, right M1 in photographic and 3D model occlusal (C) and 491 detailed occlusal (K) views; BRS 27/21, left m1 in photographic and 3D model buccal views (D); MSF 97 (BRS 27/4), 492 right hemimandible with p2 and m1 in lateral (E), occlusal (F) and detailed occlusal (L) views. Mellivora benfieldi: G- 493 H, J - MSF 88 (BRS 25/19), left hemimandible with p3-m1 in photographic and 3D model lateral (G), occlusal (H) and 494 detailed occlusal (J) views. 495 Figure 2 Map of the occurrence of the small felids of the Old World. Localities are ordered in chronological order. 496 Leptofelis vallesiensis: MN10 localities, (1) Batallones 1, (2) Batallones 3, Spain. Felis attica: MN11 localities, (3) 497 Dorn Dürkheim 1, Germany; (4) Middle Maragheh, Iran; MN12 localities, (5) Akkaşdağı, (6) Karain 2, Turkey; (7) 498 Upper Maragheh, Iran; (8) Molayan, Afghanistan, (9) Pikermi, Greece; (10) Taraklia, Moldavia; (11) Thermopigi, 499 Greece; (12) Valdecebro 5, Spain; (13) Morskaya 2; MN13 localities, (16) Las Casiones, Spain; (17) Dzedzvtakhevi, 500 Georgia. Felis christoli: MN13 localities, (14) Cava Monticino, Italy; (15) Venta del Moro, Spain; MN14 locality: (18) 501 Montpellier sands, France. Felis silvestris: (19) Ahl al Oughlam, Morocco (earliest F. silvestris); (20) Olivola, Italy 502 (type locality of F. silvestris lunensis). Data from Ficcarelli and Torre (1975); Koufos (2011); Salesa et al. (2012, 2019). 503 504 Figure 3 Biplot of the lower carnassial variables in extant and fossil small-/medium-sized felids and in the specimen 505 from Cava Monticino. 506 507 Figure 4 Carnivoran materials from Cava Monticino (Brisighella, RA, Italy). Plioviverrops faventinus: A-B – MSF 62, 508 fossiliferous block with several cranial and postcranial elements of two or more individuals in lateral (A) and detailed 509 photographic and 3D model views (B). Lycyaena cf. chaeretis: C-D – MSF 84 (BRS 27/17), cranium in right 510 photographic and 3D model lateral (C) and ventral (D) views. 511 512 Figure 5 Carnivoran materials from Cava Monticino (Brisighella, RA, Italy). Plioviverrops faventinus: A-B – MSF 92 513 (BRS 5/34), right hemimandible with p3-m2 in lateral photographic view (A); MSF 92.1 (BRS 5/34), left hemimandible 514 with p4-m2 in 3D model occlusal view (B). Lycyaena cf. chaeretis: C-F – MSF 87 (BRS 27/28), right m1 in 515 photographic buccal (C) and in 3D model occlusal (E) views; D, F, MSF 86 (BRS 27/2), right p4 in buccal view (D) 516 and in 3D model occlusal (F) views. 517 518 Figure 6 Log-ratio diagram on log-transformed dental variables of Mio-Pliocene species of Plioviverrops (namely 519 Plioviverrops gaudryi, Plioviverrops gervaisi, Plioviverrops guerini, Plioviverrops orbignyi and Plioviverrops cf. 520 orbignyi) in comparison to Plioviverrops faventinus used as a standard comparison. 521 522 Figure 7 Map of the occurrence of Plioviverrops in the Miocene of Europe. Localities are ordered in chronological 523 order. Plioviverrops gervaisi: MN5, (1) Vieux Collonges, France; MN6, (2) Calatayud, Spain. Plioviverrops gaudryi: 524 MN7/8, (3-4) La Grive 1-2, France. Plioviverrops orbignyi: MN11, (5) Ravin des Zouaves 5, Greece; (11) Gorna 525 Sushitsa, Bulgaria; MN12 localities, (12) Los Aljezares, Spain; (13) Pikermi, (14) Samos, (15) Vathylakkos-2, (16) 526 Perivolaki, (17) Prochoma, Greece; (18) Kalimantsi 2, (19) Kalimantsi 4, Bulgaria. Plioviverrops guerini: MN11, (6) 527 Piera, (7) Crevillente 2, (8) Puente Minero, (9) Viveros de Pinos, (10) Los Aguanaces, Spain; MN12, (20) Cerro de la 528 Garita, Spain; MN 13, (21) Los Mansuetos, (22) Concud, (24) La Alberca, (25) Las Casiones, Spain. Plioviverrops 529 faventinus: MN13, (23) Cava Monticino, Italy; MN14, (26) La Gloria 4, Spain. Plioviverrops cf. orbignyi: MN15, (27) 530 Çalta-1, Turkey. Data from Werdelin and Solounias (1991); Turner et al. (2008). 531 532 Figure 8 Biplot of the lower carnassial variables of selected fossil hyaenids as compared to the specimen from Cava 533 Monticino. 534 535 Figure 9 Map of the occurrence of Lycyaena in the Miocene of Europe. Localities are ordered in chronological order. 536 Lycyaena crusafonti: MN9, (1) Bled Douarah, Loc. 17, France. Lycyaena chaeretis, MN10, (3) La Roma 2, Spain; 537 MN11, (4) Hidiraga, Turkey; MN12, (5) Pikermi, (11) Samos, Greece; (12) Taraklia, Moldavia; (13) Valdecebro 5, 538 Spain; MN13, (15) El Arquillo 1, (16) Los Mansuetos, Spain. Lycyaena aff. chaeretis: MN10, (2) Los Valles de 539 Fuentidueña, Spain. Lycyaena cf. chaeretis: MN10, (10) Cerro de la Garita, Spain; (17) Cava Monticino, Italy. 540 Lycyaena macrostoma: MN12, (6) Dhok Patan, (7) Hasnot, (8) Jabi, (9) Wadia, Pakistan. Lycyaena dubia: late MN12- 541 MN13, Pao-Te-Lok.49, Wenquan-dakusitai, China. Data from Werdelin and Solounias (1991); Turner et al. (2008). 542 543 Figure 10 Biplot of the lower carnassial variables of Mio-Pleistocene Eucyon species as compared to the sample of E. 544 monticinensis from Cava Monticino. 545 546 Figure 11 Map of the occurrence of Eucyon in the Miocene-Early Pleistocene times of Europe. Localities are ordered in 547 chronological order. Eucyon cipio: MN 13, (1) Los Mansuetos, (2) Concud, Spain. Eucyon monticinensis: MN13, (3) 548 Cava Monticino, (4) Casino, (6) Verduno, Italy. Eucyon debonisi: MN13, (5) Venta del Moro, Spain. ‘Eucyon’ 549 khoikhoi: MN13, (7) Langebaanweg, South Africa. Eucyon davisi: MN13, (8) Khirgis Nur 2 Loc. 5-37, Mongolia; 550 MN14, (10) Khirgis Nur 2 Loc. 51-55, (11) Sangin Dalay Nur, Mongolia; (17) Gusinyi Perelet, Kazakhstan; (19) 551 Olkhon 520 level B, Russia; MN15-16a, (27) Xiakou, (28) Liujiagou, China. Eucyon intrepidus: MN13, (9) Lukeino, 552 Kenya. Eucyon kuta: MN 15, (21) Aralee Issie, Ethiopia. Eucyon marinae: MN15, (22) Muhor-Erig, Mongolia. Eucyon 553 adoxus: MN14-15, (18) La Calera, Spain; (23) St. Estève, France; (24) Red Crag, UK. Eucyon odessanus: MN14, (13) 554 Beresti, Romania; (14) Etulia, (15) Nikolskoe, (16) Novaja Karbolia, Moldavia; MN15, (20) Odessa Catacombs, 555 Ukraine; (25) Malusteni, Romania; (26) Megalo Emvolon, Greece; MN 17, (34) Sarikoltepe, Turkey. Eucyon zhoui: 556 MN14, (12) Gaozhuang, MN16, (29) Yinjiao, (30) Zhaozhuang, China. Eucyon minor: MN16a, (31) Shamar, (32) 557 Beregovaya, (33) Udunga, Russia; MNQ18, (36) Xiashagou, China. ‘Eucyon’ kuruksaensis: MN17, (35) Kuruksay, 558 Tajikistan. Data from Rook (2009); Valenciano et al. (2021). 559 560 Figure 12 Map of the known occurrences of Mellivora benfieldi in the Old World in comparison to the extant 561 distribution of the extant Mellivora capensis (shaded area). (1) Langebaanweg, South Africa; (2) Cava Monticino, Italy; 562 (3) Middle Awash, Ethiopia. Data from Valenciano and Govender (2020).