REVIEW OF URSUS MATERIAL FROM FONTANA RANUCCIO (MIDDLE PLEISTOCENE, CENTRAL ITALY): NEW INSIGHTS ON THE FIRST OCCURRENCE OF THE BROWN BEAR IN ITALY. Jacopo Conti1,2, Luca Bellucci3,4, Dawid A. Iurino2, Flavia Strani 2,4 , Raffaele Sardella 2,4 1 Polo museale, Sapienza Università di Roma, Italy. 2 Dipartimento di Scienze della Terra, PaleoFactory, Sapienza Università di Roma, Italy. 3 Museo di Geologia e Paleontologia, Sistema Museale di Ateneo, Università degli Studi di Firenze, Italy. 4 Istituto Italiano di Paleontologia Umana, Roma, Italy. Corresponding author: J. Conti ABSTRACT: The Middle Pleistocene archaeo-palaeontological site of Fontana Ranuccio (Anagni, central Italy) yielded a rich fau- nal assemblage consisting of more than 20,000 fossil remains, including four human teeth and multiple bone and lithic tools. The Ursus specimens from Fontana Ranuccio were historically ascribed to Ursus deningeri and Ursus arctos although a formal study has never been carried out. In this work, we have described for the first time all the Ursus material from Fontana Ranuccio reas- sessing the taxonomical attributions and discuss the first occurrence of the brown bear in the Italian Peninsula. Biometrical and morphological analyses confirm the presence of Ursus deningeri but, due the scarcity of diagnostic features and/or their preserva- tion, some isolated specimens have been ascribed to Ursus sp. The results of this work allow to reconsider the earliest occurrence of U. arctos, attested in the Italian Peninsula to the late Middle Pleistocene site of Bucine. Keywords: Ursus arctos, Ursus deningeri, Galerian, biochronology, systematics, evolution, carnivores. Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335 Alpine and Mediterranean Quaternary, 34 (1), 2021, 55-68 1. INTRODUCTION In the Middle Pleistocene locality of Fontana Ra- nuccio (0.4 Ma; Pereira et al., 2018), Ursus fossils are represented by some isolated teeth and a third phalanx, found during the 1980s excavation campaigns led by researchers of the Istituto Italiano di Paleontologia Uma- na (hereinafter IsIPU). These remains were firstly as- signed to Ursus deningeri by Biddittu et al. (1979) and Cassoli & Segre Naldini (1993) which described only three specimens without providing a detailed compari- son. Few years later, Azzaroli (1983) reported the pres- ence of the brown bear Ursus arctos in the “Ranuccio local fauna” without providing any description of the fossil remains, thus opening the issue on the possible co-occurrence of two bear species in the Fontana Ra- nuccio site. Successively, Palombo et al. (2002 and references therein) reported in their faunal list only the presence of U. deningeri at Fontana Ranuccio, but with- out performing any direct study of the fossil material. In later publications, some authors reported the first occur- rence (FO) of U. arctos at Fontana Ranuccio without supporting this statement with any comparison or analy- sis (Capasso Barbato et al., 1990; Di Stefano et al., 1994; Sardella et al., 2006; Petronio et al., 2011). Final- ly, in a recent publication, Petronio et al. (2019) consid- ered the bear remains from Fontana Ranuccio as U. deningeri, moreover indicating for this site the last occur- rence (LO) of the species. As in the previous works, the attribution was probably made on the basis of literature data, since there is a lack of direct descriptions and analyses of the materials. As a consequence of these ambiguous attributions, the taxonomy of Ursus materials from Fontana Ranuccio has been doubtful. Therefore, on the basis of the above-mentioned literature as well as from a biochronological point of view, the FO of the brown bear or the LO of the Deninger bear in Italy, or even both, could potentially be documented at Fontana Ranuccio. During the Pleistocene several fossil bear species are recorded in the Italian Peninsula, such as the Au- vergne bear (Ursus minimus, Devèze de Chabrol & Bouillet, 1827), the Etruscan bear (Ursus etruscus, Cuvi- er, 1823), the Deninger bear (Ursus deningeri, von Reichenau, 1904), the cave bear (Ursus spelaeus, Rosenmüller, 1794) and the brown bear (Ursus arctos, Linnaeus, 1758) (Conti, 1954; Conti, 2019; Borselli et al., 1980; Bon et al., 1991; Mazza & Rustioni, 1994; Palombo et al., 2002; Petronio et al., 2007, 2020; Pe- trucci & Sardella, 2009). The first occurrence of U. deningeri in the Italian Peninsula is recorded in the late Early Pleistocene sites https://doi.org/10.26382/AMQ.2021.14 56 Conti J. et al. Fig. 1 - Geographical location of the Middle Pleistocene site of Fontana Ranuccio (modified by Strani et al., 2018a). of Slivia (Ambrosetti et al., 1979; Palombo et al., 2002 and reference therein; Petronio et al., 2011, 2019). Other localities of the Early and Middle Pleistocene bearing U. deningeri remains are Bristie I° (Trentino Alto Adige; Lugli & Sala, 2000), Ponte Molle (Lazio; Petronio et al., 2011; Mecozzi et al., 2021), Selva Vecchia (Veneto; Bon et al., 1991), Iser- nia La Pineta (Molise; Peretto & Sala 2019), Visogliano (Friuli Vene- zia Giulia; Palombo et al., 2002 and reference therein) and Venosa (Palombo et al., 2002 and reference the- rein). The oldest documented and figured specimen attributed to Ursus arctos in the Italian Peninsula comes from the late Middle Pleistocene site of Bucine (Upper Valdarno, Tuscany) (Masini et al., 1991; Ferretti, 1997; Gliozzi et al., 1997; Palombo et al., 2002). The material consists in a fragmented cranium with the presence of the upper fourth premolar and the first and second up- per molars, now stored at the “Museo di Geologia e Paleontologia” at the Università di Firenze. The brown bear is also reported in Sicily from two late Middle Pleis- tocene/Late Pleistocene localities, Acquedolci and Con- trada Camillà, both referred to the “Elephas mnaidriensis faunal complex” (Bonfiglio et al., 2001; Pavia, 2001; Marra, 2003). The material is abundant but fragmented, composed by three fragmentary hemi- mandibles, isolated teeth and fragmentary postcranial bones. Other fragmented bone elements (one indeter- mined tooth and a phalanx) come from the layer 18 Ripardo del Poggio (Salerno), dated approximately to MIS 6 (Boscato et al., 2009). In this study, we present the first detailed descrip- tion and comparative analysis of the bear specimens from Fontana Ranuccio in order to: (1) define their taxo- nomic status; (2) verify the possible co-occurrence of U. arctos and U. deningeri at the site; (3) define the bio- chronological aspects as the possible FO of U. arctos or the LO of U. deningeri in the Italian Peninsula. 2. GEOGRAPHICAL AND STRATIGRAPHICAL BACKGROUND The archaeo-palaeontological site of Fontana Ra- nuccio (Fig. 1) is located in the Anagni Basin (Biddittu et al., 1979; Cassoli & Segre Naldini, 1993; Florindo et al., 2021 and references therein), about 90 km southeast of Rome, and it was discovered in 1976 during quarry ac- tivities for the exploitation of “pozzolana” (volcanic ash). Since then, researchers of the IsIPU have carried out different field activities, in particular under the direction of A. Segre (1978-2002), F. Parenti (2004-2018) and S. Grimaldi (2019 - present). The fossiliferous layer has been recently dated to 0.4 Ma (Pereira et al., 2018). The vertebrate collection includes more than 20,000 specimens that are now stored in the IsIPU depository (Anagni, Frosinone). The large mammal assemblage have been attributed to 15 taxa (Biddittu et al., 1979; Cassoli & Segre Naldini, 1993; Strani et al., 2018a; Strani et al., 2018b; Strani et al., 2019): Palaeoloxodon antiquus, Stephanorhinus sp., Equus cf. mosbachensis, Hippopotamus amphibius, Dama clactoniana, Cervus elaphus eostephanoceros, Praemegaceros sp., Bos primigenius, Sus scrofa ferus, Panthera sp., Crocuta crocuta, Canis mosbachensis, cf. Vulpes sp., Macaca sylvanus, Ursus arctos, Ursus deningeri and Ursus sp. Small mammal remains are scanty and have been recently ascribed by Bona & Stra- ni (2021) to Microtus (Terricola) sp., Talpa sp., Elyomys sp., Lepus sp., Erinaceus sp., Rodentia indet., cf. Glis sp. Finally, Homo remains have been reviewed by Rubi- ni et al. (2014) and are associated to several (more than 800) lithic and bone tools (Segre, 1984). Since the pio- neering work of Azzaroli (1983) this faunal assemblage is a reference for the Italian biochronological framework (Gliozzi et al., 1997; Petronio et al., 2011; Petronio et al., 2019 and references therein). Finally, Grimaldi et al. (2020) carried out a techno-functional study of the entire lithic assemblage identifying five groups of retouched tools i.e., cutting tools, pointed tools, scrapers, notches and denticulates. 3. MATERIALS AND METHODS Ursus material from Fontana Ranuccio consists of 12 isolated remains (Fig. 2) now stored at the IsIPU laboratory in Anagni (Frosinone). This fossil material was mainly found in the 1980s field activities, in particu- lar in the 1982, 1984, 1985 and 1989 years. Morphologi- cal analysis, descriptions and comparisons have been provided taking into account the from different research- ers (Ballesio, 1983; Torres, 1988; Capasso Barbato et al., 1990; Argant, 1991; Mazza & Rustioni, 1994; Augu- ste, 1995; Rabeder, 1999; Quiles, 2003; Meloro, 2007; Wagner & Sabol, 2007; Wagner & Čermák, 2012). Mor- phological data have been compared with specimens of U. etruscus, U. deningeri, U. spelaeus and U. arctos 57 Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy) Tab. 1 - List of selected specimens used for morphological comparison. Asterisks indicate the material directly studied by the authors. from selected European Pleistocene and Holocene lo- calities (Tab. 1). The ontogenetic stage was inferred according the scheme proposed by Stiner (1998). The measurements of mesiodistal length and buc- colingual width have been taken with digital caliper to the nearest 0.1 mm on each fossil specimen (Tab. 2), with the exception of the teeth FR 85-1 S1, FR84-1 and the third phalanx (FR 84 inv 56576) due fragmentary preservation. Dentognathic morphology and measure- ments follow the terminology proposed by Rabeder (1999) and Torres (1988). Despite the high biometric variability of the teeth in the Early and Middle Pleistocene bears, we performed standard bivariate plots of width vs. length of M1, p4, m2 and m3 in order to compare our samples with speci- mens from this time span in order to test possible bio- metrical patterns among the Fontana Ranuccio speci- mens and other Pleistocene ursid taxa (Torres, 1984; Capasso Barbato et al., 1990; Marra, 2003; Rabeder et al., 2010; Wagner et al., 2017 and personal measure- ments; Fig. 3). 4. SYSTEMATIC PALAEONTOLOGY Class Mammalia Linnaeus, 1758 Order Carnivora Bowdich, 1821 Family Ursidae Fisher [De Waldheim], 1817 Genus Ursus Linnaeus, 1758 Ursus sp. Referred material: I2 right (FR Sd-2), i3 right (FR 85-2 P3), i3 left (FR 82-2 Res), c right (FR 85-1 S1), p4 right (FR 89-1), m1 left (FR 96-33), m2 left (FR 84-1), ph -3 (FR 84 Inv 56576). Incisors and fragmented material: the incisors, except FR Sd-2, are well preserved and show the typical Ursus morphology with a developed central cusp and the presence of two outlined lateral lobes; the distal one is less developed than the mesial one (Fig. 2 f-g-h). The lower canine (FR 85-1 S1), the second lower molar (FR 84-1) and the first lower molar (FR 96-33) are fragment- ed; therefore, it is not possible to acquire any linear measurement or identify any distinctive character to define a specific attribution (Fig. 2 i-l-m). However, it is possible to ascribe the specimen FR 96-33 to a prime adult individual (class VII). The third phalanx (FR 84 inv 56576, Fig. 2 n) shows a rather elongated morphology, although it is particularly worn. The proximal portion, which is articulated with the trochlea of the second phal- anx, is semicircular and the dorsal and plantar swelling that characterize this anatomical portion of the bone element are poorly developed. The ungual portion is thin with a gentle dorsal curvature until the most distal por- tion, where it sharply curves. Due to high wear and the lack of any discriminant characters, we ascribe these remains to Ursus sp. Lower fourth premolar: the lower fourth premolar (FR 89-1) has a very simple morphology, with narrow and elongated external profile. The protoconid is well developed and undivided, unlike the metaconid and the paraconid which are small and poorly developed. The presence of a tubercled area in the occlusal surface is not observed except for one small accessory cusp on 58 Conti J. et al. T a b . 2 - L is t a n d M e a s u re s o f U rs u s m a te ri a l fr o m t h e F o n ta n a R a n u c c io d e p o s it ( M e a s u re s i n m m ). Quiles, 2003; Madurell-Malapeira et al., 2009; Rabeder et al., 2010; Wagner & Čermák, 2012; Arsuaga et al., 2014; Prat-Vericat et al., 2020). This simpler morphology is related to the early Pleistocene Etruscan bear U. etruscus, characterized by the lack of accessory cusps or tubercled structures in the chewing surface of molariforms, and considered as ancestor of both lineages (Wagner & Sabol, 2007). The teeth of the Late Pleistocene cave bear (Ursus ex gr. spelaeus) show instead a higher complexity of the chew- ing surface (Fig. 4 k-l), contrary to what is observed in the Late Pleistocene and extant brown bears, which possess a simpler and secodont morphology of the oc- clusal surface through time (Fig. 4 m-n-o-p). These dif- ferences are probably related with the dietary habits of the two bear lineages, one omnivorous (U. arctos) and the other mostly herbivorous (U. spelaeus) (Terlato et al., 2018). Biometric analyses did not allow any specific discrimination regarding the Middle Pleistocene speci- mens (Fig. 3 b), while the dimensional range of U. the lingual portion of the protoconid slope. All these fea- tures occur in the arctoid tooth morphology and can be found also in the Late Pleistocene and recent Ursus arctos (Reynolds, 1906; Torres, 1988; Capasso Barbato et al., 1990; Mazza & Rustioni, 1994; Quiles, 2003; Me- loro, 2007; Rabeder et al., 2010). U. deningeri can dis- play a higher morphological complexity of the occlusal surface characterized by many tubercles and by a much more elliptic external profile (Fig. 4 e-f-g-i), even if many remains such as the early Middle Pleistocene speci- mens from the site of Hundsheim (Austria, HH 305), C718 Cave (Czech Republic, Rv 20003) (Fig. 4 d-h), and some other specimens recorded from Sima de los Huesos (Spain, Middle Pleistocene, ca. 0.35 Ma), Caune de l’Arago, (France, Middle Pleistocene, ca. 0.50 Ma), Cal Guardiola (Spain, latest Early Pleistocene, ca. 1.2-0.86 Ma), display a p4 without any accessory cus- pids and simpler morphology, remarking the high varia- bility and the possible presence of ancestral characters displayed by U. deningeri (García et al., 1997, 2007; 59 Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy) Fig. 2 - Ursus material from Fontana Ranuccio in occlusal and lateral views: a) M1 right (FR 06-411), b) FR M1 left (FR 56574), c) p4 right (FR 89-1), d) m2 left (FR sd-1), e) m3 right (FR 56575), f) i3 left (FR 82-2 Res), g) i3 right (FR 85-2 P3), h) I2 right (FR Sd-2), i) c right (FR 85-1 S1), l) m2 left (FR 84-1), m) m1 left (FR 96-33), n) III phalanx right (FR 84 inv 56576). 60 Conti J. et al. Fig. 3 - Comparison of the length and width scatter plots of various Early and Middle Pleistocene species of the genus Ursus. a) M1; b) p4; c) m2; d) m3. deningeri and U. arctos strongly overlap (also due to the lack of sufficient data of the Early/Middle Pleistocene Ursus arctos). As mentioned above, a simple morpholo- gy, with the lack of accessory cusps in the lower p4 is a shared character between Ursus arctos and Ursus deningeri, specially during the late Early-Middle Pleisto- cene (Wagner et al., 2017). Despite the presence of Ursus deningeri in the deposit, the large variability of the morphology and biometric data do not allow a specific attribution of the isolated p4. To be more conservative we ascribe FR 98-1 to Ursus sp. Ursus deningeri von Reichenau, 1904 Referred material: M1 left (FR 56574), M1 right (FR 06-411), m2 left (FR sd-1), m3 right (FR 56575). Upper first molars: the specimen FR 56574 (Fig. 2 b) shows sub-quadrangular profile with a non-rectilinear protocone-hypocone line and the presence of some tubercled structures on the talon surface. The central constriction of the tooth is rather marked especially in the lingual portion and the main cusps are well devel- oped. The biometric data show, as expected, an overlap between the deningeroid and arctoid specimens; despite this, we can observe that FR 56574 falls within the di- mensional range mostly occupied by the Early-Middle Pleistocene U. deningeri (Fig. 3 a) and showing a rather large size of this tooth compared to the average dimen- sion of U. arctos. The specimen FR 06-411 (Fig. 2 a) has some dif- ferent features from FR 56574. The metastyle and the parastyle are well developed, the metastyle more than the parastyle, and the talon shows a tubercled surface, more than FR 56574. On the other hand, the tooth has a quadrangular shape, the distal and proximal area show the same profile and the protocone-hypocone line ap- pears quite straight. Biometric analyses indicate that the specimen is placed at the limit of the average dimen- sional range of U. arctos, falling within the variability of U. deningeri (Fig. 3 a). The occlusal surface of both teeth displays tubercles, a common speloid feature (Capasso Barbato et al., 1990; Quiles, 2003). According to Torres (1984) and Capasso Barbato et al. (1990) the straight protocone-hypocone line of FR 06-411 should be related to the arctoid lineage. The latter assumption stands up against the morphological comparison here proposed (Fig. 5), in fact specimens of U. deningeri from Hundsheim (Fig. 5 e-f) show the same straight proto- ipoconid line, more than brown bear, which instead pre- sents a consistent variability for this character. Consid- ered all these morphological and biometric features, we ascribe these specimens to U. deningeri. Second lower molar: The specimen FR sd-1 (Fig. 2 d) is characterized by a high complexity of the occlusal surface with the presence of numerous tubercled struc- tures and accessory cusps, well representing the typical cave bear morphology (Fig. 6) consisting of a metaconid d i v i d e d i n t o t h r e e metastylids, typical of U. deningeri and U. ex gr. spe- laeus (Capasso Barbato et al., 1990; Quiles, 2003; Torres, 1988). The biometric analysis still indicates a straight dimensional overlap between U. deningeri and U. arctos, which does not allow to distinguish the two species based on size. The specimen FR SD-1 falls perfectly in this overlap range (Fig. 3 c). Con- cerning the morphological features of the m2 from Fon- tana Ranuccio we ascribe this specimen to U. deningeri. Lower third molar: The specimen FR 56575 shows a quadrangular outline with a rounded talonid. The occlusal surface is highly tubercled, typical of the speloid lineage (Capasso Barbato et al., 1990; Quiles, 2003; Torres, 1988) (Fig. 7). The tooth shows low worn structures and a well-developed ento- conid and metaconid, the latter is composed by two distinct cusps. Biometric data also suggest the affinity of the specimen with the speloid group, fitting within the dimensional range of U. deningeri (Fig. 3 d). Accord- ing to both morphological and biometric analyses, we ascribe this specimen to U. deningeri. 5. DISCUSSION AND FINAL REMARKS The presence of U. deningeri is confirmed at Fontana Ranuccio, as al- ready proposed by Biddittu et al., (1979) and Cassoli & Segre Naldini, (1993). This species is widely distributed throughout Europe and quite well represented in Italy during the Middle Pleistocene (Fig. 8). U. deningeri was a medium-large sized bear generally deemed as the ancestor of Ursus spelaeus s.l. The transition between these two species occurred during the Middle - Late Pleistocene transition, so far these taxa are considered two chronospecies, phylogenetically closely related (Sardella et al., 2006, Valdiosera et al., 2007). From molecular studies a complex scenario emerges during the Late Pleistocene, with three different evolutionary lines: U. spelaeus, U. ingressus and U. deningeri kudarensis (Knapp et al., 2009). Late Pleistocene cave bear in particular retained the same character of the Middle Pleistocene Deninger bear, even if more marked, 61 Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy) Fig. 4 - Comparison of the p4: a) Ursus arctos (left), DA-4B 18-36 (Deutsch-Altenburg, Early Pleisto- cene); b) Ursus etruscus (right, mirrored in the table), IGF 911 (Valdarno, Early Pleistocene); c) Ursus etruscus (left), IGF 4605 (Valdarno, Early Pleistocene); d) Ursus deningeri (left), HH 305 (Hundsheim, early Middle Pleistocene); e) Ursus deningeri (left), HH 306 (Hundsheim, early Middle Pleistocene); f) Ursus deningeri (left), Rv 20003 (Koněprusy Caves, early Middle Pleistocene); g) Ursus deningeri (left), Rv 20005 (Koněprusy Caves, early Midde Pleistocene); h) Ursus deningeri (left), n.c. (Isernia la Pineta, early Middle Pleistocene); i) Ursus deningeri (left), specimen H (Sandalja, Middle Pleistocene); j) FR 89-1 (right, mirrored in the table) (Fontana Ranuccio, Middle Pleistocene); k) Ursus savini nor- dostensis ssp. (left), IAM F-2365 (Ovrag, Late Pleistocene); l) Ursus spelaeus (left), P3021 (Caverna delle Fate, Late Pleistocene); m) Ursus arctos (left), ZIN 34595 (Kudaro 3, Late Pleistocene); n) Ursus arctos (left), V.1152 (Vigna S. Carlo, Late Pleistocene); o) Ursus arctos (left), 3399 (Alps, Recent); p) Ursus arctos (right, mirrored in the table), 3362 (Alps, Recent). Abbreviations: Med Metaconid, Prd Protoconid, Pad Paraconid. as the broad, domed, steep forehead, that result in a ‘step’ in the midsagittal plane (Santos et al., 2017). During the Middle Pleistocene, U. arctos has been reported in some Italian localities: Bucine (Upper Val- darno, Tuscany; Mazza, 1998), Riparo del Poggio (Salerno; Boscato et al., 2009), Acquedolci and Contra- da Camillà (Sicily; Bonfiglio et al., 2001; Marra, 2003; Pavia, 2001) (Fig. 8). The brown bear record from Ri- paro del Poggio is composed only by two elements (one indeterminate tooth and a phalanx), the deposit consists in a shelter which was part of a complex underground karst system, which was partially dismantled by sea erosion. The Ursus arctos material come from the lower level dated to the cold phase of MIS 6 (Boscato et al., 2009). The cranium from Bucine comes from one of many small fossiliferous deposits as Poggio Amaro, Le Capannelle, Pogi di Bucine, Capannole, Migliorini, Cava di Bucine, Cava Le Vigne, Cava del Rinoceronte, mostly situated along the Ambra creek, a left-hand tributary of the Arno river, and excavated since the 1940. No infor- mation is available on the exact locality where the crani- um of U. arctos was recovered, but all the fossil remains are dated around the late Middle Pleistocene (referable to the Saalian cycle, from 340 ky to 120 ky ago; Mazza, 62 Conti J. et al. Fig. 5 - Comparison of the M1. a) FR 06-411 (right). b) FR 56574 (left, mirrored in the table). c) U. deningeri (left, mirrored in the table), HH/5/342 (Hundsheim, Middle Pleistocene) d) U. deningeri (left, mirrored in the table), HH/5/398, (Hundsheim, Middle Pleistocene). e) U. arctos (left, mirrored in the table), DA4B/18 (Deutsch-Altenburg, Early Pleistocene). f) U. arctos (right), DA4V/14 (Deutsch-Altenburg, Early Pleistocene). g) U. arctos priscus (right), W S.1 (Winden, Late Pleistocene). h) U. arctos (left, mirrored in the table), C. 4 (Banskà Bystrica, Re- cent). i) U. arctos (right), IGF 10961 (Bucine, late Middle Pleis- tocene). e-f-g-c) modified from Rabeder et al. (2010). Fig. 7 - Comparison of the m3. a) FR 56575 (right). b) U. arctos (left, mirrored in the table), C.4, (Banská Bystrica, recent). c) U. deningeri (left, mirrored in the table), MF/1346/45 (Kozi Grzbiet, Middle Pleistocene). d) U. deningeri (left, mirrored in the table), Hund 382 (Hundsheim, Middle Pleistocene). Fig. 6 - Comparison of the m2. a) FR 84-1 (left). b) FR sd-1 (left). c) U. arctos (right, mirrored in the table), 3364 (Alps, recent). d) U. deningeri (right, mirrored in the table), 1889/5/404 (Hundsheim, Middle Pleistocene). e) U. deningeri (left), MF/1346/37 (Kozi Grzbied, Middle Pleistocene), modified from Wagner et al. (2012). 1998). The bear remains from Sicily have been excavated in the last forty years, and have been found associat- ed with Cervus elaphus siciliae and Elephas mnaidriensis, dated to the late Middle Pleistocene/Late Pleistocene (E. mnaidriensis Faunal Complex, from early MIS 6 to MIS 4 stages; Bonfiglio et al., 2001, 2003). The occurrence of the brown bear is also documented in some other sites of the north/eastern Italian Peninsula (see Bon et al., 1991), however this material needs further chronological and systematic analyses. Since the uncertain dating of these deposits, it is possible to hypoth- esize that the Bucine specimen could be the F.O. of the brown bear in the Italian peninsula. A recent paleoenvi- ronmental reconstruction based on dietary adaptations of ungulates from Fontana Ranuccio suggests that a mosaic of both closed and open land- scapes characterized this locality around 0.4 Ma with a relative abun- dance of soft plant resources and probably affected by a marked season- ality (Strani et al., 2018a; Strani et al., 2019; Strani, 2020; Strani et al., 2021). A heterogeneous environment could have favored the coexistence of two different bear species in the area, which may have lived in sympatric conditions exploiting different trophic resources as largely documented in several Late Pleistocene localities of the Italian Peninsula (Capasso Barbato et al., 1990; Minieri et al., 1995; Pa- lombo et al., 2002; Mazza et al., 2005). Although palaeoecological con- siderations allow the presence of the two species, paleontological data do not show with any reliable evidence of the presence of Ursus arctos. The fossils from Fontana Ranuccio repre- sent the LO of Ursus deningeri in Italy. New excavations are still ongoing car- ried on by IsIPU and new findings will better address many unsolved ques- tions and shed new light on the distri- bution and evolution of bears in Europe. ACKNOWLEDGEMENTS We are very grateful to the Italian Institute of Hu- man Paleontology for granting access to the material and for the continuous assistance during the analyses. We would like to express also gratitude to Martin Sabol (Comenius University, Bratislava), for advising and the inspiring discussion that made possible this work. We also thank the MuSe (Museo delle Scienze of Trento), the Parco Nazionale d’Abruzzo, Lazio e Molise, the Department of Geology and Paleontology of Comenious University (Bratislava, Slovakia), the Slovak Museum of Nature Protection and Speleology (Liptovský Mikuláš, Slovakia), the National History Museum (Wien, Austria), the Department of Paleontology of Wien University (Wien, Austria), for granting access to fossil and extant bear material used for morphological comparisons. We thank Chiara Delpino of Soprintendenza Archeologia Belle Arti e Paesaggio per le Province di Frosinone, Latina e Rieti (Ministero della Cultura) and the Italian Institute of Human Paleontology for granting access to 63 Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy) Fig. 8 - Distribution of the fossil bears during the Middle Pleistocene of the Italian Peninsu- la. 1) Venosa (van Heteren et al., 2019). 2) Bristie I (Lugli & Sala, 2000). 3) Cava Rinaldi (Petronio et al., 2019). 4) Cretone (Petronio et al., 2011). 5) Fontana Ranuccio (this work). 6) Isernia la Pineta (Peretto & Sala, 2019). 7) Ponte Molle (Petronio et al., 2011). 8) Slivia (Bon et al., 1991; Petronio et al., 2011). 9) Valdemino (Ghezzo et al., 2015). 10) Visoglia- no (Tozzi et al., 2000). 11) Cava Nord (Soave) (Rustioni & Mazza, 1993). 12) Cengelle 1 (Soave) (Rustioni & Mazza, 1993). 13) Castello (Soave) (Rustioni & Mazza, 1993). 14) Cerè (Ghezzo et al., 2013; Rossi & Santi, 2011). 15) Selva Vecchia (van Heteren et al., 2019). 16) Sorbano di Romagnano (Bon et al., 1991). 17) Fornace di Cornedo (Rustioni & Mazza, 1993). 18) Grotta di San Bernardino (Bon et al., 1991). 19) Acquedolci (Marra, 2003). 20) Contrada Camilla (Marra, 2003). 21) Bucine (Mazza, 1997; Petronio et al., 2019). 22) Castel di Guido (Caloi et al., 1998). 23) Prati Fiscali (Petronio et al., 2011). 24) Fara Sabina (Petronio et al., 2011). 25) Riparo del Poggio (Boscato et al., 2009). 64 Conti J. et al. the palaeontological collections of Fontana Ranuccio.". 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