A NEW HYENA DEN FROM THE LATE PLEISTOCENE BUCA DELLA JENA CAVE (ROSELLE, GR): PRELIMINARY REPORT Omar Cirilli, Saverio Bartolini Lucenti, Andrea Savorelli, Carlo Cavanna, Andrea Sforzi & Luca Pandolfi O. Cirilli, Dottorato di Ricerca in Scienze della Terra, Università di Pisa, Via S. Maria 53, 56126, Pisa, Italy. Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121, Firenze; omar.cirilli@phd.unipi.it S. Bartolini Lucenti, Dottorato di Ricerca in Scienze della Terra, Università di Pisa, Via S. Maria 53, 56126, Pisa, Italy. Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121, Firenze; saverio.bartolini@dst.unipi.it A. Savorelli, Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121, Firenze; andrea_savorelli@yahoo.it C. Cavanna. Società Naturalistica Speleologica maremmana, Museo di Storia Naturale della Maremma, Grosseto, Strada Corsini 5, 58100, Grosseto; carlocavanna1@gmail.com A. Sforzi. Museo di Storia Naturale della Maremma, Grosseto, Strada Corsini 5, 58100, Grosseto. L. Pandolfi. Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121, Firenze; luca.pandolfi@unifi.it KEY WORDS - Hyena den, Late Pleistocene, Mammals, Maremma, Tuscany, Italy INTRODUCTION Numerous Late Pleistocene paleontological sites in the Italian peninsula have yielded fossil assemblages that come from hypogeal contexts. Most of these sites are known and well documented in literature since the beginning of 20th century. As far as Southern Tuscany is concerned, caves are particularly abundant in the Colline Metallifere area and in the Natural park of Maremma (Guerrini, 1985; Cavanna, 1998). In 1997, the discovery of a new cave from some members of the Società Naturalistica Speleologica Maremmana has unveiled one of the richest fossil cave assemblages of the area. The cavity, since then known as “Buca della Jena” is located near the town of Roselle (Grosseto, Figure 1) on the southern slope of Poggio di Moscona, a calcareous hill entirely composed of lower Jurassic limestone, 200 metres above sea level. Buca della Jena is part of a large karst network as many others karstic cavities are known in Poggio Moscona area (Cavanna, 1998). Our preliminary results show that this mammal assemblage is largely dominated by herbivores, in particular Equidae. Confirming the first interpretation given by the speleologists who named the cave, the presence of several coprolites favours the idea that the deposit originated from a hyena den, albeit a taphonomic analysis is still needed for a definitive confirmation of this hypothesis. Undergoing pollen analyses are expected to give insights on the environment at the time of accumulation. The Buca della Jena mammal assemblage is noteworthy since it represents one of the richest Late Pleistocene fossil assemblages from Southern Tuscany together with Grotta Rose Mary (early Late Pleistocene), Grotta di Golino (MIS4-3) and Grotta La Fabbrica (MIS4-3) (cfr. Petronio et al., 2011). SYSTEMATIC NOTES Crocuta crocuta Erxleben, 1777 Figure 2, A. The spotted hyena appeared in Italy during Middle Pleistocene times (Ponte Galeria Faunal Unit; Sardella & Petrucci, 2012) and its last record is during the MIS 3 (Stuart & Lister, 2014). The species was widely distributed across Europe and well adapted to a great variety of habitats and climatic conditions (Conti et al., 2012). The fossil of Crocuta sample includes few fragmented cranial dentognatic remains (among which a neurocranium), isolated teeth, and several well preserved coprolites (22 specimens). The specimens belong to adult individuals, as no deciduous tooth or unfused bones were recovered in the sample. Another evidence of the presence of the spotted hyena is the prominent bite and gnawing marks on several bones in the assemblage. Ursus arctos Linnaeus, 1758 Figure 2, B. The first occurrence of Ursus arctos in Italy is usually placed during the second half of the Middle Pleistocene (Caloi & Palombo, 1997) or at the end of the Middle Pleistocene (Petronio et al., 2011). Remains of this species are more abundant in Late Pleistocene localities, sometimes in association with Ursus spelaeus (e.g., Monte Cucco Cave, Breuil Cave) (Petronio et al., 2011). Ursid remains are relatively abundant in the studied Cave. They are represented by fragments of skull and mandible, isolated teeth, and several postcranial remains which seem to belong to the same individual (both left and right bones are documented and are similar in size). The morphology and morphometry of teeth and bones fit well with an attribution to the brown bear Ursus arctos. 1 Canis lupus Linnaeus, 1758 Figure 2, C-D. The earliest record of the modern wolf in Italy is that of La Polledrara di Cecanibbio (Anzidei et al., 2012). After that, Canis lupus becomes a common element of late Middle Pleistocene and Late Pleistocene deposits, regardless the climatic conditions (Sansalone et al., 2015; Mecozzi & Bartolini Lucenti, 2018), managing to survive until today. In the sample from Buca della Jena, the modern wolf is represented by few isolated dental specimens (e.g., left P4 and a right m1), and by a single cranial specimen (preserving only the part of the neurocranium). Some morphological feature (e.g., P4 general morphology, development of the P4 protocone, and presence of metaconid in the m1) exclude the attribution of the specimens to Cuon alpinus (Pallas, 1811). No peculiar features allow to differentiate these fossils from the extant wolf, therefore the specimens are referred to the species Canis lupus. Figure 2, C-D. Vulpes vulpes Linnaeus, 1758 Figure 2, E-F. The fossil record of foxes is more elusive than other carnivorans. Among the first record of the modern red fox, there is that of L’Escale (Bonifay, 1971), while in Italy is reported in La Polledrara di Cecanibbio (Anzidei et al., 2012). In the Buca della Jena, the red fox is represented by a fragmented mandible and isolated dental specimens. The morphology and development of the cusps and cuspids are consistent with those of Vulpes vulpes, ruling out a possible attribution of the sample (or part of it) to the smaller and more carnivorous arctic fox Vulpes lagopus. Equus ferus Boddaert, 1785 Figure 2, G. Equus ferus may has spread in Europe in the Middle Pleistocene from central Asia, and gave rise to a relevant number of subspecies, considered different from several authors by sizes proportions (e.g. E. ferus germanicus, E. ferus gallicus, E. ferus gmelimi). The earliest record of the wild horse Equus ferus in Italy is within the Isernia Faunal Unit, middle Galerian. The morphology of wild horses is different from the other Late Villafranchian stenonoid horses in the structure of the upper teeth enamel, and in the morphology of the protocone (more asymmetrical than in stenonoid horses), as well as in the postcranial remains where the metapodials are thicker and more compressed antero–posteriorly (Conti et al., 2010); first and second phalanges of the central digit, instead, are more developed in dimensions (length and width). In the Buca della Jena mammal assemblage Equus is represented by 48 specimens, among which 35 are cranial remains (upper and lower isolated teeth, one fragmentary left upper teeth row, one fragmentary left mandible), and 13 postcranial specimens. The upper and lower teeth show the typical caballine occlusal morphology (Eisenmann 1980; 1981), with an asymmetrical protocone in upper teeth and a pli-caballin on the hypoconid in the lower teeth. Postcranial specimens have measurements and proportions coherent with the Pleistocene wild horse E. ferus. In the Buca della Jena mammal assemblage the fossil horses are represented by Equus ferus. Remarkable is that all the fossil horses specimens are from E. ferus; no remains of the small Late Pleistocene Equidae Equus hydruntinus have been found, contrary to other fossil assemblage in the central and South of Italy (Conti et al., 2010). Rupicapra rupicapra Linnaeus, 1758 Figure 2, H. Rupicapra rupicapra probably occurred in Italy at the end of the Middle Pleistocene (Masini, 1985), but it is relatively well-documented during the Late Pleistocene, in particular it is recorded in fossiliferous localities referred to the Last Glacial period (Petronio et al., 2007). At present, the species is distributed in mountain areas of southern Europe and Near East but during the Pleistocene it also occurred in low-altitude areas and coastal area (Massetti & Salari, 2012). Among the Buca della Jena material, a fragmented small mandible with hypsodont teeth and a calcaneum are assigned to the chamaois. Cervus elaphus Linnaeus, 1758 Figure 2, I. C. elaphus occurred for the first time in Italy at the Early–Middle Pleistocene transition, with the subspecies C. elaphus acoronatus. During the Middle Pleistocene several other subspecies (i.e. C. elaphus eostephanoceros, C. elaphus rianensis and C. elaphus aretinus) are documented in central Italy. Populations of red deer with antler morphology similar to that of the extant subspecies occurred in the Peninsula at the beginning of the Late Pleistocene (Di Stefano & Petronio 1993, 2002; Di Stefano et al. 2015). Although variations in body size of C. elaphus have been reported by several scholars, recent studies suggested that red deer populations are characterised by stasis in body size evolution, with a non-significant interaction with climate and a significant relationship with geography (Pandolfi et al., 2015). The red deer is particularly common in open woodland with large grassy clearings (Mattioli, 2003). Cervus elaphus is represented by numerous remains, including fragments of skulls and mandibles, upper and lower tooth rows and postcranial remains. The morphology of the collected specimens resembles that 2 of the extant C. elaphus and the morphometric values are close to the maximal values reported from this species (data collected from Di Stefano et al., 2015; Pandolfi et al., 2015). Arvicola amphibius Linnaeus, 1758 This large and semi-aquatic arvicolid appears in Italy at the beginning of the Late Pleistocene (Sala & Masini, 2007) and is a common element in mainland Italy during Late Pleistocene and Holocene (Gatta et al. 2018). A. amphibius is typical of waterside habitats and an indicator of locally humid conditions (Masini & Abbazzi, 1997; Benvenuti et al., 2017). The water vole is well represented in the assemblage, its remains consist of 20 specimens. Among these, 7 mandibles still bearing cheek teeth, 4 edentulous mandibles and a fragmental maxillary without teeth, and 8 isolated teeth. The molars are apparently characterized by an undifferentiated dental enamel (sensu Heinrich, 1978, see also Maul et al. 2014). Microtus (Microtus) arvalis Pallas, 1778 The common vole is represented by 14 specimens: a mandible still preserving m1 and m2, a fragmental palate bearing both M1s, M2s and right M3, and 12 isolated teeth. The diagnostic dental elements such as m1s show the typical morphology of M. arvalis (Chaline et al. 1974), whereas non diagnostic teeth can be reasonably attributed to the same species due to the comparable size and morphology. The presence of this arvicolid indicates cool climatic conditions. In fact, nowadays Italian populations of M. arvalis are limited to the northern regions of the peninsula (Toschi, 1965; Benvenuti et al., 2017). Lepus sp. The hare occurs frequently in Middle and Late Pleistocene Italian sites (Kotsakis et al., 2003). The material attributed to the hare consists of 34 isolated teeth (6 upper incisors, 9 lower incisors and 19 cheek teeth), 2 fragmental maxillaries (one still preserving M1), 4 fragmental mandibles (3 of which still preserve some teeth). The size of the remains seems to exclude an attribution to the genus Oryctolagus. DISCUSSION In the Italian Peninsula the Late Pleistocene was characterized by several climatic events that influenced the faunal composition, with latitudinal migrations and the contraction of the suitable viable areas for several species (Petronio et al., 2011). At the beginning of the Late Pleistocene, some taxa recorded since the Middle Pleistocene were still present (e.g., Palaeoloxodon antiquus, Stephanorhinus hemitoechus, and, rarely Hippopotamus amphibius); Bos primigenius and E. ferus were relatively abundant. Amongst the Cervidae, the red deer occurred with subspecies characterized by the antler morphology of extant Cervus elaphus ssp., and the extant fallow deer (Dama dama dama) occurred for the first time (Di Stefano & Petronio, 2000; Petronio et al., 2007). The cold stages of the Late Pleistocene were characterized by the presence of Coelodonta antiquitatis (which probably occurred for the first time in Italy during the Middle Pleistocene; Pandolfi & Marra, 2015), Mammuthus primigenius and Marmota primigenia. During these time spans Stephanorhinus, Palaeoloxodon and Hippopotamus gradually disappeared from the Peninsula (Petronio et al. 2007; Pandolfi & Petronio, 2016; Pandolfi et al., 2017), whereas Capra ibex and the chamois expanded their areas of distribution (Petronio et al. 2007). Among carnivores, C. crocuta was well-documented during the Late Pleistocene and the species became extinct in the Peninsula during late MIS 3 (around 31 ka cal. BP: see Stuart & Lister 2014). M. arvalis, typical of open environments and rather cool climate, was widespread throughout the Peninsula but the species was never recorded during MIS 5 and the Late Holocene in central and southern Italy (Kotsakis et al., 2003 with references, 2011; Paolucci & Amori, 2008; Tang & Kotsakis, 2008; López García et al., 2014, 2018; Salari, 2014). A. amphibius was documented in almost all Late Pleistocene and Holocene fossiliferous sites of mainland Italy (Kotsakis et al., 2003 with references, 2011). Accordingly, the fossiliferous locality of Buca della Jena can be referred to a relatively cool stage of the Late Pleistocene, older than ca. 31 ka cal. BP (last occurrence of C. crocuta in Italy). CONCLUSIONS The locality of Buca della Jena represents a database of precious information for MIS 3 faunas of Southern Tuscany due to the scarcity of fossiliferous sites in that area. Indeed, only other two sites from the same area can be referred to this time span: Grotta di Golino and Grotta La Fabbrica. The presence of hyena coprolites and the large number of gnaw-marks allowed us to reasonably interpret the deposit as a hyena den. Nevertheless, a deeper taphonomic analysis of the deposit and an age-profile structure of the collected mammals is needed to 3 confirm this hypothesis, as well as to reconstruct the accumulation’s history of the deposit and the past landscape of southern Tuscany in a time span characterised by important changes in climates and faunal compositions. ACKNOWLEGEMENTS The present authors thank the members of the Società Naturalistica Speleologica Maremmana for the discovery of the cavity and the documentation of its fossil content, F. Landucci (Vertebrate Paleontology Laboratory at the Earth Science Department of Florence) for technical support, E. 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New radiocarbon evidence on the extirpation of the spotted hyena (Crocuta crocuta (Exrl.) in norther Eurasia. Quaternary Science Review, 96: 108-116. TANG Z. & KOTSAKIS T. (2008). Tardiglacial micromammals of Riparo Salvini (Latium, Central Italy). Geologica Romana, 41: 125-132. TOSCHI A. (1965). Mammalia. Lagomorpha, Rodentia, Carnivora, Artiodactyla, Cetacea. Fauna d’Italia, Vol. VII, Calderini, Bologna, 647 p. Figure 1. Geographical locator map of the Buca della Jena cave, Roselle (GR). 5 Figure 2. Buca della Jena (Roselle, Grosseto) Late Pleistocene: Crocuta crocuta - (A) BJ123: left hemimandible fragment. Ursus arctos - (B) BJ124: left hemimandible fragment. Canis lupus: (C) BJ041: right m1; (D) BJ099_e: right P4. Vulpes vulpes: (E) BJ044: left m1; (F) BJ090: right hemimandible fragment. Equus ferus - (G) BJ096: left maxillary fragment. Rupicapra rupicapra - (H): BJ050: left hemimandible fragment. Cervus elaphus - (I): BJ035: right hemimandible fragment. All pictures are in labial view. Scale bar 5 cm. 6