AMQ104 Mecozzi et al 3 compresso.pub Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335 Alpine and Mediterranean Quaternary, 33 (1), 2020, 89-98 CUON ALPINUS (PALLAS, 1811) FROM THE LATE PLEISTOCENE SITE OF INGARANO (FOGGIA, SOUTHERN ITALY) AND INSIGHTS ON THE EURASIAN MIDDLE TO LATE PLEISTOCENE RECORD. Beniamino Mecozzi 1, 2, Saverio Bartolini Lucenti 3, 4, Dawid Adam Iurino 2 1 Dipartimento di Scienze della Terra, Università di Roma Sapienza, Roma, Italy. 2 PaleoFactory, Università di Roma Sapienza, Roma, Italy. 3 Dipartimento di Scienze della Terra, Paleo[Fab]Lab, Università degli Studi di Firenze, Firenze, IItaly. 4 Museo di Storia Naturale, sez. Geologia e Paleontologia, Università degli Studi di Firenze, Firenze, Italy. Corresponding author: Saverio Bartolini Lucenti ABSTRACT: In this study, we report for the first time the presence of Cuon alpinus from the Late Pleistocene site of Ingarano (Foggia, southern Italy), represented by an right upper first molar. Considering the intricate and debated taxonomy of fossil dholes, our comparative analyses on dental samples (P4, M1, and M1) of the extant and Middle to Late Pleistocene dholes from Europe, has been performed evidencing a relevant degree of morphological variability and a biometric uniformity of the considered teeth. Our results indicate the lack of clear morphological and biometric features for a reliable teeth-based classification of fossil dholes, questioning the validity of the fossil taxa currently proposed in the literature. Finally, to avoid the propagation of taxonomically questionable species, we suggest to refer all the Middle to Late Pleistocene material to Cuon alpinus. Keywords: Canidae, carnivorans, dhole, biochronology, biometry, teeth.   1. INTRODUCTION Cuon alpinus (Pallas, 1811), known as dhole, is a medium-sized social canid currently widespread in most of southeastern Asia including the western islands of Indonesia (Durbin et al., 2004; Wilson & Mittermeier, 2009; Castelló, 2018). Morphological and biometric data led Mivart (1890) to distinguish two extant species of Cuon, the northern dhole (C. alpinus) and the southern dhole (C. javanicus). On the contrary, Ellerman & Morri- son-Scott (1966) reunited the diversity of the dhole un- der a single specific name, identifying 11 subspecies on the basis of their external features (e.g., differences in coat length, color pattern) (Castelló, 2018). Genetic analyses confirmed Ellerman and Morrison-Scott’ hy- pothesis, finding no clear specific distinction, although such works only took into consideration populations from the southern part of the dhole distribution (e.g., Iyengar et al. 2005). Wilson & Mittermeier (2009) grouped all these subspecies under only three, C. al- pinus alpinus, C. alpinus hesperius, C. alpinus suma- trensis. Therefore, there is a general consensus in con- sidering the genus Cuon as monospecific, although the validity of the different subspecies is still debated. The dhole is an active predator that lives in large packs, characterized by several dental features related to a hypercarnivorous diet (Van Valkenburg, 1991), such as the slicing premolars, the single-cusped talonid of the lower carnassials, the reduction or even the ab- sence of the second upper molar and the lack of the lower third molar. Except for this latter character, peculi- ar of the genus Cuon, the hypercarnivorous dentition is shared with the extant genera Lycaon and Speothos (Thenius, 1954; Van Valkenburg, 1991) and the fossil Xenocyon. Whereas today the species is confined to Asia, at the beginning of the Middle Pleistocene the dhole was widespread across Eurasia reaching western Europe (García García, 2003) and North America in the Late Pleistocene (Tedford et al., 2009). Such a wide geo- graphical distribution is a common ecological trait of the Middle-Late Pleistocene canids as documented for Ca- nis mosbachensis and Canis lupus (Sardella et. al., 2014; Bartolini Lucenti et al., 2017; 2020; Mecozzi et. al., 2017, 2020), both characterized by a large intraspe- cific variability and thus considered as polymorphic spe- cies. Fossil remains of dhole are exceptionally scarce and mainly represented by isolated teeth, fragmented cranial remains and few postcranial elements (Perez- Ripoll et al., 2010). Consequently, the morphological and biometric variability of these carnivorans throughout the Middle and Late Pleistocene is poorly known, fueling the debate on its origin and evolution (Brugal & Boudadi- Maligne, 2011, Petrucci et al., 2012; Ghezzo & Rook, 2014). Here, we offer a comparative study of selected teeth (P4, M1 and M1) of extant and fossil dholes, includ- ing the well preserved right M1 belonging to the first Cuon alpinus reported from the Late Pleistocene site of Ingarano. https://doi.org/10.26382/AMQ.2020.08 90 Mecozzi B. et al. 1.1. Taxonomy of the European fossil dholes In the last quarter of the XIX Century, in Europe several taxa have been ascribed to the genus Cuon (C. alpinus fossilis, C. alpinus pyrenaicus, C. bourreti, C. europaeus) (Tab. 1). Thenius (1954) erected the spe- cies Cuon priscus on the basis of the large-sized speci- mens from Hundsheim (Austria). Furthermore, the au- thor introduced the subspecies C. dubius stehlini for the fossils from Rosières (France), suggesting similarities with the Chinese C. dubius described by Teilhard de Chardin (1940). Nowadays, the species “C. dubius” is commonly referred to the genus Xenocyon (Schütt, 1973; Tedford et al., 2009) or even the transitional Sin- icuon (Wang et al., 2014). The form “C.” stehlini is now referred to Lycaon/Xenocyon lycaonoides (Rook, 1994; Martínez-Navarro & Rook, 2003, Sotnikova & Rook, 2010). Similarly, the species Cuon rosi from Cueva Vic- toria described by Pons-Moyá & Moyá-Solá (1978) is widely considered as Lycaon/Xenocyon lycaonoides (Madurell-Malapeira et al., 2013). The reason for this confusion between the Pleistocene remains of Europe- an dholes and those of genus Lycaon/Xenocyon lies in the similar development of several dental features. Ad- am (1959) attempted to reconstruct the affinities and taxonomy of European Cuon, suggesting the division of the European remains in three subspecies, all part of a single phyletic line: C. a. priscus - C. a. fossilis - C. a. europaeus. Bonifay (1971) considered valid three spe- cies for the Middle Pleistocene of Europe: C. stehlini, C. priscus and C. alpinus (the latter furtherly diversified into two subspecies, C. alpinus fossilis and C. alpinus euro- paeus). The subspecies C. alpinus caucasicus was erected by Baryshnikov (1978) on the basis of the Late Pleistocene fossil remains from various localities of the Transcaucasian region (Kudaro 1, Kudaro 3, Tsona cave). Recently, Brugal & Boudadi-Maligne (2011) re- vised the scheme proposed by Bonifay (1971) acknowl- edging three taxa for the Pleistocene fossil record of Europe: C. priscus, C. a. fossilis and C. a. europaeus. As mentioned above, the first occurrence of C. priscus has been reported from the early Middle Pleistocene site of Hundsheim (Austria) (Thenius,1954). Afterwards, this species has been documented until the Middle Pleisto- cene in some localities e.g., Lunel-Viel (France) (Bonifay, 1971), Caune de l’Arago (France) (Bonifay, 1971), Ga- leria Pesada (Portugal) (Trinkaus et al., 2003). During the late Middle Pleistocene, the first occurrence of anoth- er species of dhole, C. alpinus, has been reported from the late Middle Pleistocene site of Trinchera Galeria in Spain (Garcìa Garcìa & Arsuaga, 1997; Garcìa Garcìa, 2003). This taxon probably replaced C. priscus becoming part of the carnivoran guild for all the Late Pleistocene Tab. 1 - List of the fossil and extant (sub)species of Cuon. * indicates extant subspecies listed in Wilson & Mittermeier (2009). Fossil spe- cies are marked by † symbol. Fig. 1 - Geographic position of the Ingarano: a) map of Italy; b) zoom of the promontory of Gargano (image taken from GoogleEarth). 91 until the Early Holocene, as reported from the Riparo Fredian site (Italy) (Cilli et al., 1998) and Les Coves de Santa Maira (Spain) (Pérez-Ripoll et al., 2010). 1.3. The site of Ingarano The palaeontological site of Ingarano (Foggia, southern Italy) is located in the north-western area of the Gargano’s promontory (Fig. 1). The deposit is a karstic filling succession within the Jurassic-Cretaceous limestone (Petronio et al., 1996; Petronio & Sardella, 1998; Bedetti & Pavia, 2007) dated with the 238U/234Th method (Tab. 2). A number of studies focused on the description of the rich vertebrate fauna of Ingarano site (levels B-E, Fig. 2) (Capasso Barbato et al., 1992; Petronio et al., 1996; Petronio & Sardella, 1998; Curcio et al., 2005; Bedetti & Pavia, 2007; Iurino, 2014; Iurino & Sardella, 2015; Iurino et al., 2015), including 41 species, listed below: Amphibians: Rana sp.; Reptiles: Lacerta sp.; Birds: Circus aeroginosus, Aquila chrysaëtos, Falco cherrug, Falco columbarius, Alectoris graeca, Perdix perdix, Columba livia, Nyctea scandiaca, Pyrrhocorax graculus, Pyrrhocorax sp., Corvus corone, Corvus corax; Mammals: Erinaceus europaeus, Myotis blythi, Oryctolagus cuniculus, Lepus europaeus, Arvicolidae indet., Microtus sp., Microtus ex gr. arvalis/agrestis, Terricola savii, Apodemus sylvaticus, Eliomys querci- nus, Canis lupus, Vulpes vulpes, Ursus arctos, Mustela nivalis, Martes sp., Meles meles, Gulo gulo, Crocuta crocuta, Felis sp., Lynx lynx, Panthera pardus, Equus hydruntinus, Cervus elaphus, Dama dama, Capreolus capreolus, Rupicapra sp., Bos primigenius. The faunal assemblage was deposited in a short time-span, chrono- logically referred to MIS 3 (Bedetti & Pavia, 2007). The talus, a deposit of unclear stratigraphical interpretation found near the sedimentary succession, is rich in verte- brate remains, including Panthera spelaea, Hippopota- mus amphibius, Coelodonta antiquitatis, Stephanorhinus hemitoechus and Palaeoloxodon antiquus. No fossil remains of these taxa have been recovered from the levels B-E. Up to now, the remains of medium-sized canid have been attributed to Canis lupus (Petronio et al., 1996; Petronio & Sardella, 1998; Bedetti & Pavia, 2007). 2. MATERIALS AND METHODS The specimen INGND59 representes a right upper first molar and it is currently housed in the PaleoFactory laboratory, Department of Earth Sciences, at the Sapien- za University of Rome. Comparative fossil samples include: specimens of San Sidero, housed in the PaleoFactory laboratory, De- partment of Earth Sciences, Sapienza University of Rome; those of “Grotta” and “Tecchia di Equi”, housed in the collection of the Geology and Paleontology section of the Natural History Museum of the University of Flor- ence; and specimens of Wanxian housed in the Verte- brate Paleontology collections of the American Museum of Natural History (AMNH, New York, U.S.A.). The taxon from Wanxian was previously referred to C. antiquus by Matthew & Granger (1923) but currently is considered a junior synonym C. alpinus, see among others Castelló Fig. 2 - Fossiliferous site of Ingarano: a) schematic log of the stratigraphic succession; b) sedimentary succession. Tab. 2 - Radiocarbon dating of speleothem and phosphathic material from the site of Ingarano (modified from Petronio et al., 1996). Cuon Alpinus (Pallas, 1811) from Ingarano (Foggia, southern Italy) and insights on the Eurasian Middle to Late Pleistocene record 92 Mecozzi B. et al. Fig. 3 - Variability of M1 in extant and fossil Cuon. a–d) Cuon alpinus from Ingarano, INGND59, right M1; a) buccal view; b) lingual view; c) occlusal view; d) schematic outline of the occlusal morphology of INGND59. e–g) extant Cuon alpinus; e) AMNH 102083; f) IOZ 26747 (Tong et al., 2012); g) Cuon alpinus hesperius Afanasjev & Zolotarev, 1935, holotype, ZIN 23894, Zoological Institute of Russian Academy of Science. h–l) fossil Cuon. h) Cuon alpinus, AMNH 18391 (reversed), Wanxian (China); i) Cuon alpinus, AMNH 18727 (reversed), Wan- xian (China); j) Cuon alpinus europaeus Bourguignat 1875, Obarreta (Portugal, from Perez-Ripoll et al., 2010); k) Cuon priscus Thenius, 1954, holotype from Hundsheim (Germany, from Thenius, 1954); l) Cuon alpinus caucasicus Baryshnikov, 1978, Kudaro (from Baryshnikov, 2012). (2018) (Tab. 1). Data on the extant C. alpinus were tak- en on the specimens of the Comparative Anatomy AMNH (New York, U.S.A.). Following Driesch (1976), we measured the length (L) and breadth (B) of the P4, M1 and M1. The measure- ments were taken to the nearest 0.1 mm with a digital caliper. We used literature data on fossil dhole from the Middle to Late Pleistocene of Austria, Czech Republic, Slovakia, France, Germany, Greece, Italy, Spain, Hun- gary and Caucasian region (See supplementary materi- al, Tab. S1). The specimens from Romain La Roche, Mosbach and Petralona have not been considered for biometric and morphological analyses as we exclude their attribution to the genus Cuon (see Discussion). Following Brugal & Boudadi-Maligne (2011), we consid- er the European fossil sample belonging to two taxa, C. priscus from the early to late Middle Pleistocene and C. alpinus from the late Middle to Late Pleistocene (Tab. S1). The dataset was tested for normality distribution verification using Shapiro-Wilks test. As the dental varia- bles resulted normally distributed, we performed a sta- tistical parametric test. Differences in P4L, M1L, and M1L of the Middle to Late Pleistocene specimens of Europe was evaluated using linear model with corner point pa- rameterization testing the null hypothesis of no- differences between the mean of C. priscus and the other sample of Middle and Late Pleistocene C. alpinus and extant C. alpinus (p.value > 0.05). The statistical analysis was performed using the R software (R Core Team, 2019). 3. SYSTEMATIC PALAEONTOLOGY Family Canidae Fischer, 1817 Genus Cuon Hodgson, 1838 Cuon alpinus (Pallas, 1811) 3.1. Description INGND59 is an almost complete right M1 missing part of the labial root (Fig. 3A-C). The crown is slightly worn and slightly wrapped by a crust formed by the clay matrix with calcite cementum. In occlusal view, the M1 is buccolingually short. It possesses an enlarged paracone compared to the slightly shorter metacone. These cusps appear to be fused medially. In occlusal view, the buccal cingulum is not prominent and it is only marked at the level of the parastyle. The trigon basin is wide, round and deep, mesially girdled by a stout and large proto- 93 Cuon Alpinus (Pallas, 1811) from the Late Pleistocene site of Ingarano (Foggia, southern Italy) and insights on the Eurasian Middle to Late Pleistocene record cone. The post-protocrista distally shows a very reduced and barely visible metaconule. The talon basin is vestigi- al and a cingulum, although it is partially broken, lies lingually. The mesial cingulum is feeble. The embay- ment on the distal side of the tooth is rather reduced. 3.2. Morphological comparison The fusion of the paracone and the metacone, as well as the development of the buccal cingulum of INGND59, resemble the condition observed in: C. priscus from Hundsheim (Thenius, 1954), C. alpinus caucasicus from Caucasus (Kudaro, Baryshnikov, 1996), C. alpinus europaeus from Spain (e.g., Obarreta, Peréz-Ripoll et al., 2010) and in those from Italian area of Melpignano and San Sidero (Petrucci et al., 2012; Iurino et al., 2013) (Fig. 3). The morphology of the lin- gual portion of the M1 has a high degree of intraspecific variability (also at individual level, see Altuna, 1983). In occlusal view, the lingual portion of the INGND59 has a round and expanded outline similar to those of the spec- imens from Hundsheim (Thenius, 1954) (Fig. 3k), Wan- xian (“Wan Hsien”, Sichuan, China; Colbert & Hooijer, 1953) (Fig. 3h-i), Trinchera Galeria (García García, & Arsuaga, 1997), and different from that of other fossil and extant samples (e.g., Kudaro) (Fig. 3l) for their re- duction of the distolingual lobe. A peculiar feature of the M1 from Ingarano is the relative enlargement of the lin- gual portion compared to the buccal one. This morpholo- gy contrast with that observed in extant Cuon where generally the lingual side of the tooth is mesiodistally compressed (Fig. 3). 3.3. Biometric comparison The results of the statistical analysis reveal slight differences in the dental traits (Fig. 4). In extant dhole, the range of the P4L includes all the values of the fossil specimens, which are very similar to each other as the median of C. priscus reaches the maximum value of the extant dhole. The length and breadth of the P4 in C. priscus do not differ significantly from the other groups (p.values > 0.05) (Fig. 4a-b; Tab. 3). The Middle Pleisto- cene forms possess longer M1 compared to the other groups, but its breadth is significantly lower than those of C. priscus and the Late Pleistocene C. alpinus (Fig. 4d). Similarly, the M1B of C. priscus is larger compared to the Middle Pleistocene C. alpinus (Fig. 4c-d; Tab. 3). Whereas the M1L of C. priscus statistically differs from those of the Late Pleistocene C. alpinus and the extant C. alpinus, no differences can be found in the breadth (Fig. e-f; Tab. 3). Tab. 3 - Table of p-values of linear model, Cuon priscus used as corner point. 94 Mecozzi B. et al. Fig. 4 - Boxplot of the length (a) and breadth (b) of P4, the length (c) and breadth (d) of M1 and the length (e) and breadth (f) of M1 of Mid- dle and Late Pleistocene Cuon alpinus and extant Cuon alpinus. be the reason of the proliferation of the fossil species/ subspecies/varieties of Cuon (Brugal & Boudadi- Maligne, 2011) (Tab. 1). The tooth from Ingarano falls in this range of morphologies although its lingual portion is among the widest of the considered sample. A certain degree of teeth variability among the Italian record of fossil Cuon has been reported by Petrucci et. al. (2012), where the authors highlighted how the shape and the occlusal surface of the M2 change in all the Italian speci- mens. A similar variability of the M2 in extant and fossil dhole has been reported also by Brugal & Boudadi- Maligne (2011). According to Mivart (1890) and Durbin et al. (2004) the M2 variability documented in extant dhole would be linked to the latitudinal distribution, but for a more exhaustive explanation of such a teeth diver- sification a larger sample of fossil specimens is needed. The Eurasian fossil record of the dhole is mainly represented by isolated or fragmentary cranio-dental elements, whereas postcranial bones and partially com- plete crania are exceptionally rare, which explains why 4. DISCUSSION According to our results, the specimen INGND59 is morphologically and biometrically compatible with the M1 of extant and fossil dhole, thus representing the first report of this carnivoran from the site of Ingarano. De- spite the fact that the remains of Cuon are poorly docu- mented and both geographically and chronologically dispersed, our morphological comparison has evidenced a quite large variability of the buccal and lingual profile of the M1 among the considered samples, especially the cuspule-like postprotocrista distal to the metacone and both the buccal and distal notches (Fig. 3c-d). This vari- ability would not appear to be taxon-related as the ex- tant forms include almost all the range of features ob- served in the fossil samples, as exemplified by the three different M1 morphotypes identified in the extant dhole specimens (Fig. 3e-g). Along with other variable dental features (e.g., the development of cuspulid-like lingual portion of the M1 talonid), these subtle differences, might 95 Ms. received: July 10, 2020 Final text received: September 08, 2020 Fig. 5 - Length variation of the P4, M1 and M1 of the European fossil Cuon from the Middle Pleistocene to the present days. Middle Pleisto- cene: 1) La Caune de l'Arago, 2) Lunel Viel, 3) Hundsheim, 4) Heppenloch; Late Pleistocene: 5) Trinchera Galeria; Late Pleistocene: 6) Repolust Hole, 7) Chokier, 8) Certova dira, 9) Sipka, 10) Ceverne Mars/Vence, 11) Fontechevade, 12) Grotte de Cotencher, 13) Grotte de la Carrière, 14) Grotte de l'Observatoire, 15) Grotta Vaufrey, 16) Isturitz, 17) La Niche, 18) Malarnaud, 19) Moula Guercy, 20) Verze, 21) Offenberger Hole, 22) Melpignano, 23) San Sidero, 24) Buca del Tasso, 25) Canale Mussolini, 26) Equi, 27) Ingarano, 28) Valserra, 29) Almada, 30) Bolinkoba, 31) Boquete de Zafarraya, 32) Cova Negra, 33) Duranguesado, 34) La Riera, 35) Moros de Gabasa, 36) Obarreta, 37) Parpallò, 38) Kudaro, 39) extant. Cuon Alpinus (Pallas, 1811) from Ingarano (Foggia, southern Italy) and insights on the Eurasian Middle to Late Pleistocene record the taxonomy of this carnivoran is principally based on teeth. Moreover, biometric data as the size of the lower carnassial, were considered as good parameters to discriminate the dhole from other Canidae (Ghezzo & Rook, 2014) or to distinguish between the fossil sam- ples and to recognize different taxa (Brugal & Boudadi- Maligne, 2011). Nevertheless, up to now no statistical analyses were performed to support these hypotheses. The results of our study carried out on the P4, M1 and M1, reveal that size differences among the extant and fossil samples are very small, with weak, if any, statisti- cal significance (Fig. 4, Tab. 3). Differences can be found only in some variables without any consistent evidence of distinction between the known taxa. For instance, the M1 length of the Middle Pleistocene C. alpinus reaches the highest values of the sample whereas the M1 breadth is larger in C. priscus and in the Late Pleistocene C. alpinus. In general, the variability of the fossil groups is mostly included in that of the extant species, suggesting a biometric uniformity of the teeth size from the Middle Pleistocene to recent times. This contrasts with the hypothese put forward by Adam (1959), with three recognized forms (C. alpinus priscus, C. alpinus fossilis, C. alpinus europaeus in Adam 1959) based on the supposed trend of size decrease in the M1 length. Such idea was confirmed by Brugal & Boudadi Maligne (2011). These authors separated the Middle Pleistocene apparently larger forms (C. priscus) from the Late Pleistocene ones (C. alpinus) on the basis of teeth size. The authors included among the Middle Pleistocene forms the samples from Mosbach, Petra- lona and Hundsheim, which are indeed characterized by larger teeth (Tab. S1). Nevertheless, it should be noted that the specimen of Mosbach was considered by Adam (1959) as Cuon priscus, but this material has not been described. Schütt (1974) ascribed the Mosbach specimen to Xenocyon lycaonoides for the retention of the M3 in the hemimandible. Similarly, the left hemi- mandible from Petralona (KP No 205) was originally described by Kurtèn & Poulianos (1977) as belonging to C. priscus. The authors remark that the hemimandible is partially covered by a stalagmite crust, especially in the molar area, which prevents the acquisition of any bio- metric or morphological data of the M2 and on the pres- ence/absence of the M3. Therefore, the classification was carried out without considering these characters. More recently, Baryshnikov & Tsoukala (2010) revised the fossil collection from Petralona stored at the Aristo- tle University of Thessaloniki, identifying three canid taxa: Canis arnensis, Vulpes praeglacialis and Lycaon lycaonoides (=Xenocyon lycaonoides). In the latter, the authors also included the remains previously classified as Cuon priscus by Kurtèn & Poulianos (1977) into L. lycaonoides (Baryshnikov & Tsoukala, 2010: 392). An- other questionable attribution to Cuon is that of a large- sized specimen from Romain La Roche (CAN.1069) described by Argant (2010). The latter scholar reported the presence of the M3 alveolus, yet there is a wide consensus among researchers in considering the ab- sence of this tooth as one of the principal diagnostic characters to typify the genus Cuon from other canids (Schutt, 1973; Tedford et al., 2009). On such basis, the 96 Mecozzi B. et al. attribution of this specimen to Cuon alpinus is excluded and a more parsimonious attribution to Canis lupus could be suggested. According to this taxonomic clarifi- cation, we excluded the specimens from Mosbach, Petralona and Romain La Roche from our biometric analysis, evidencing the inconsistency of the time- related reduction of the teeth in European fossil dholes proposed by Adam (1959) and Brugal & Boudadì Mal- gine (2011). Indeed, size of the teeth seems to be uni- form from the Middle Pleistocene to the present days (Figs. 4-5; Tab. S1), questioning the taxonomic reliabil- ity of the fossil specimens currently proposed in litera- ture. In sum, our results indicate the lack of clear mor- phological and biometric features for a reliable classifi- cation of the European fossil dholes, especially if based on teeth. Therefore, to avoid the proliferation of taxo- nomically questionable species, we suggest to consider all the Middle to Late Pleistocene material as Cuon alpinus. 5. CONCLUSION The revision of the palaeontological collection from Ingarano allow us to identify for the first time the pres- ence of C. alpinus in this Late Pleistocene site, enrich- ing its carnivorans guild. This makes the Ingarano de- posit one of the richest Late Pleistocene Italian sites in carnivorans, where also the occurrence of C. lupus is documented. The results of this work show that some Pleisto- cene specimens that Cuon has been often confused with other hypercarnivorous taxa (e.g. C.lupus or Xeno- cyon spp.), thus confusing its taxonomic determination. Especially size was a feature historically used as the discrimination factor between different species/ subspecies. Our analyses point out that no significant difference really exists between Middle and Late Pleis- tocene forms. This dimensional uniformity, at least in teeth size, contrasts with the evidence on other canids with a rather more conspicuous record (e.g., C. mosbachensis or C. lupus, among others Sansalone et al., 2015; Mecozzi et al., 2020) which experienced sev- eral shifts in size during glacial/interglacial phases (Mecozzi & Bartolini Lucenti, 2018). If this could be re- lated to the bias of the record (i.e. its scarcity) or to the paleodietary and paleoecological aspects of these canids, are among the questions that still need to be answered. In the meantime, considering 1) the present knowledge of the fossil Cuon, 2) its scarce record in the Middle-Late Pleistocene of Europe, 3) its extant variabil- ity in morphology; our results favour the inclusion of the whole Middle-Late Pleistocene fossil record under the specific name of Cuon alpinus, rather than in two or more different (chrono) species. 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